Ecological Relations of Roots

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Papers of John E. Weaver (1884-1956) Agronomy and Horticulture Department
1-1-1919
Te Ecological Relations of Roots
John E. Weaver
University of Nebraska
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Weaver, John E., "Te Ecological Relations of Roots" (1919). Papers of John E. Weaver (1884-1956). Paper 13.
htp://digitalcommons.unl.edu/agronweaver/13
THE ECOLOGICAL RELATIONS OF ROOTS
BY
JOHN E. WEAVER
fl'
Professor of Plant Ecology in the University of N roraska
PUBLISHED BY THE CARNEGIE INSTITUTION OF WASHINGTON
WASmNGTON, 1919
CARNEGIE INSTITUTION OF WASHINGTON
PUBLICATION No. 286
PRESS OF GIBSON BROTHERS
WASHINGTON. D. C.
CONTENTS.
PAGE PAGE
List of mustrations..... . . .. ......... T I Introduction.... . ..... . .... ... . ....... 1
I. THE PRAIRIES.
Panicum virgatum.......... ........... 4 Brauneria pallida...................... 11
Andropogon furcatus....... . ...... .... . 4 Petalostemon candidus.... . . .... .... ... 11
Andropogon scoparius...... . . . .. . . .. . . . Vernonia. baldwinii .................. :.. 12
Andropogon nutans.................... 5 Kuhnia glutinosa...................... 12
Stipa spartea........ .. ....... .... ... . . 6 Verbena stricta....... . . .. .. .... .. ..... 12
Kceleria cristata..... . . .. . .. .... . . . . . . . 6 Grindelia squarrosa.... . .. .. .. . . . . . . ... 12
Elymus canadensis........ . . ........ . . . 6 Glycyrrhiza lepidota... ... .. .. .. . . .. . .. 13
6 Astragalus crassicarpus................. 13
DlBtlchlis splcata............. . ....... . 6 Psoralea tenuiflora......... . . . .. . ... . .. 14
Sporoboluslongifolius .. . J..,':'; C. . . .. .. . 7 Psoralea argophylla..... .. .. . . .. . . . .... 14
Aristida oligantha. . . . . . . . . . . . . . . . . . . . . 8 Baptisia bracteata.... . . . . . . . . . . . . . . . . . 14
Bulbilis dactyloides. . . . .. . . . . . . . . . . . . . . 8 The Subclimaz Prairie. . .. . .. . . . . . . . . . . 15
Bouteloua gracilis......... . . .... . . . . . . . 8 Brauneria pallida ............ , . . .. . . . .. 15
9 Lygodesmia juncea............. ....... 16
Liatns scanosa ...... , ..... , . . . ... . .. .. 9 Lespedeza capitata......... . . . .... . . . . . 16
Solidago rigida ........... , ., .. . . . . . . . 9 Ceanothus ovatua......... .. .. . . . . . . . . . 17
Solidago canadensis.................... 11 Amorpha canescens.................... 17
Silphium laciniatum........... . . .. . .... 11 Prairie Root SY8te7n8 and Prairie Environ-
Amorpha canescens.................... 11 ment......................... 18
Helianthus rigidus.... .. .. .... .. .. . . ... 11
II. THE CHAPARRAL COMMUNITY.
Symphoricarpos vulgaris......... . .. .... 25 Vitia vulpina...... . .. . . . . .. .. . . . . . . ... 27
Rhus glabra........................... 26 Rosa arkansana....................... 27
Corylu8 americana. . .. . . . . . . . . . . . . . . . 27 Shrub Root SY8t67n8 and the Shrub En-
vironment.. . . . . . . . . . . .. . . . . . . . 28
Ill. THE PRAIRIES OF THE PACIFIC NORTHWEST.
The Root Systems 01 the Gra8BfJ8.......... 31 Heuchera glabella........... .. .. ....... 37
Agropyrum spicatum................... 31 Leptotrenia multifida.......... ......... 37
Festuca ovina ingrata ... : ....... . . . . ... 33 Helianthella douglasii .... , . . . . . . . . . . . .. 37
Poa sandbergii.. . .... . . .. . . . ... . . . . . .. 33 Hoorebekia racemosa......... . . . . . . . . .. 37
Kceleria cristata....................... 33 ruderale.... ..... .. .. ..... 37
The Root Systems 01 other Prairie Species. 33 Sleversla ciliata. . . . . . . . . . . . . . . . . . . . . . .. 37
Lupinus omatus...... .... .. ...... . .. 33 Sidalcea oregana........ . . . . . . . . . . . . . . 37
Lupinus leucophyllus............ . ... ... 35 Hieracium scouleri....... . .. . . . .. . .. . .. 38
Astragalus arrectus..... .. . . . . . . .... . .. 35 Potentilla blaschkeana.......... .. . .. ... 38
... 35 Eriogonum heracleoides................ 38
Geramum VlSCOSISSlmum........ ........ 35 Prairie Root SystemB and Prairie Environ-
Wyethia amplexicaulis................. 35 ment... ............. ... ...... 38
IV. THE PLAINS ASSOCIATION.
Bouteloua gracilis. . . . . . . . . . . . . . . . . . . .. 46 Lygodesmia juncea. . . . . . . . . . . . . . . . . . .. 53
Aristida purpurea... . .. . . . . . . . . .. . . . ... 46 Aragallus lambertii ................... , 55
Muhlenbergia gracillima...... ... . . . . . .. 47 Petaloatemon purpureus..... . . . . . . . . . .. 55
Gutierrezia sarothrl8.......... . . . ... . .. 49 Petalostemon candidus.... .. . . . . . . . . . .. 57
Psoralea tenuiflora...... . .. . .. .. . . . . . .. 49 Eriogonum jamesii....... .. . .. . . .. . . . .. 57
Artemisia frigida. . .. . . . .. . . .. . . .. .. ... 50 Ratibida columnaris. .. . .. . . . . . . . . . . .. . 59
Argemone platyceras...... . .. . ......... 50 Senecio aureus oblanceolatua ...... " . . .. 59
Yucca glauca........ . . .. . . .. . . . . . . . .. in Asclepias verticillata pumila... . . . . . . . .. 59
Agropyrum .. s.'n1Y:4.j........ 52 Opuntia camanchica................... 61
Carel!: pennsylvanica...... .... ...... ... 52 Opuntia fragilis......... .. . .. . .... .. . .. 62
Andropogon scoparius...... ... .... . . ... 52 Plains Root SYBte7n8 and the Plains En-
Stipa comata........... .... .... ....... 53 vironment..... . .. . . . . .. . .. . ... 63
Lithospermum linearifolium. . . . . . . . . . . .. 53
nI
IV CONTENTS.
V. THE SANDmLLS SUBCLnIAX.
PAGII
fiexuosa. . . . . . . . . . . . . . . . . . . .. 68
Calamovilfa longifolia ................. , 68
Andropogon hallii. . . . . . . . . . . . . . . . . . . . .. 70
M uhlenbergia pungens. . . . . . . . . . . . . . . .. 70
Sporobolus cryptandrus. . . . . . . . . . . . . . .. 71
Eriogonum microthecum. . . . . . . . . . . . . . .. 71
Artemisia filifolia. . . . . . . . . . . . . . . . . . . . .. 73
Tradescantia virginiana. . . . . . . . . . . . . . . . 73
PAGII
Heliotropium convolvulaceum. . . . . . . . . . . 74
Petalostemon villosus.................. 74
Gilia longifiora........... ............ 75
Euphorbia petaloidea. . . . . . . . . . . . . . . . .. 76
Psoralea lanceolata. . . . . . . . . . . . . . . . . . . . 78
lpomrea leptophylla. .. . . . . . . . . . . . . . . . . 78
SandhiU Roo' Sf/ltema aM the SandhiU
Environment. . . . . . . . . . . . . . . . 79
VI. THE COMJroNITY.
81 Thlaspi alpestre....................... 86
Paronychia jamesn......... ............ 82 Mentzelia multiflora................... 87
Aletes acaulis......................... 83 Eriogonum ftavum... ................. 87
Apocynum androBlllmifolium .... , ..... ,. 84 Grovel-Slide Rooe Sf/lItema and Gravel-Slide
Smilacina stellata......... .... .... . . ... 85 Environment. ............... 88
Pachylophus cmspitosus................ 85
VII. THE HALF-GRAVEL-SLIDE COMJroNITY.
Elymus triticoides... . . . . . . . .. .. . . ..... 92 Gilia aggregata........... ........... .. 95
Solidago oreophila. . . . . . . . . . . . . . . . . . . . . 92 Potentilla arguta glandulosa. . . . . . . . . . .. 97
Rubus deliciosus........ . . . ... .. . . .. . .. 93 FraBera speciosa.......... ............. 97
Besseya plantaginea......... . . ... . . . ... 94 Aster porteri....... ............... ... 98
Geranium cleSpitosum.... .... . . .. .. ... 95 Hall-Gro:/Iel-Slide Root Sf/ltema and the
Calamagrostis purpurasceus....... . . .... 95 Half-Gra'lJel-Slide Environment... 98
Kreleria cristata. . . . . . . . . . . . . . . . . . . . . .. 95
VIII. THE FOREST COMMUNITY.
Pirola chlorantha ........ , ............. 100 Rosa acicularis ....................... 104
Thalictrum fendleri .................... 100 Arctostaphylos uva-ursi ............... 105
Erigeron asper .......................... 101 Senecio cernuus ....................... 105
Erigeron macranthus ................... 101 CastiJleia miniata ................ 105
Fragaria virginiana ... , ... " ........... 102 Heuchera parvifolia ................. 105
Allium cernuum ............... , ....... 103 Saxifraga bronchialis ................. 106
Aralia nudicaulis ........... , .......... 103 Haplopappus PIUTYi .. 107
Opulaster opulifoliuB. . . . . . . . . . . . . . . . . .. 103 Forest Root Sf/ltema and the Forest lOS
Ribes lacustre. . . . . . . . . . . . . . . . . . . . . . . .. 104 Environment.
IX. ECADs.
Smilacina stellata ...................... 110
Chammnerium angustifolium. . . . . . . . . . . .. 110
Elymus triticoides. . . . . . . . . . . . . . . . . . . .. 112
Bouteloua gracilis. . . . . . . . . . . . . . . . . . . . .. 112
Stipa comata ......... , ................ 113
Yucca glauca .......................... 114
Allionia linearis; ..... . . . . . . . . . . . . . . 114
Abronia fragrans .................. 115
Kreleria cristata....................... 117
Cbrysopsis villosa ................... 117
Euphorbia .................... 119
Root Modi/ieationa of Pol1J(lemic8 ..... ... 121
X. SUMMARY.
Summary ............................. 122 I Bibliography .................. ' ........ 128
LIST OF ILLUSTRATIONS.
PLATES.
PLA-TE A. . . . .
Quadrat-bisect showmg root distrlbutlOn
of certain dominant and BUbdominant
plains species: A, Ariatida vurvurea;
B, B01deloua gracilis; Ar, Artemisia
friqida; p, Psoralea tenuijlora; C,
ChrysopsiB villo3a; Y, Yucca glauea.
PLA-TE B.
Quadrat-bisect in the half-gravel-slide. The
face of the trench was cut along the
front of quadrat shown in Plate 25, A:
S, Solidago oreophila; A, Allium cer-
nuum; E, Elym'U8 triticoides, fragments
of which are shown in blue; C, Cala-
magrostis VUr']YUra8C1J118; H, H euchera
paruiJolia; B, Bess61/a plantaginea.
PLA-TE C.
Quadrat-bisect showing root systems of
shrubs and herbs of the forest floor.
This was made along the front edge
of the quadrat shown in Plate 27, B:
PLA-TE 1.
H, Haplopappm parrvi; R, Rosa acicu-
laria; Pi, Pin'U8 jlexilis; F, Fragaria
tnrginiana; C, Chamamerium angu8-
tifolium; P, Picea engelmanni.
A. One end of a trench used in excavating
root systems.
B. Distichlis spicata, showing the long rhi-
zomes and shallow roots.
PLATE 2.
A. A prairie of eastern Nebraska in June,
Erigeron ram08'U8 and M ericlix ser-
ru/ata in the foreground, and bushy
Psoralea ttmuiftqra in background.
B. An area dominated by Agrostis himnalis,
with AUium mutabile, Achillea mille-
folium, and Stipa spartea.
PLA-TE 3.
A. Andropogon furcatus.
B. Andropogon scopari'U8.
C. Andropogon nutans.
PI.ATE 4.
A. Panicum tnrgatum, showing rhizomes,
coarse roots, and complete single root.
B. Stipa spartea.
PLATE 5.
A. Kmleria cri8tata.
B. ElymUB canadensis.
PLATE 6.
A. SporobolUB longifoliUB.
B. B01deloua gracilis, excavated near the
quadrat shown in text-figure 1.
PI.ATE 7.
A. Silphium laciniatum, roots partially exca-
vated.
B. Amorpha canescens, showing the wide
lateral spreaa.
PLATE 8.
A. Kuhnia gllttiM8a, roots partially exca-
vated.
PLATE 8--eontinued.
B. Glycyrrhiza lepidota; a' is a continua-
tion of a.
PLATE 9.
A. A8tragai'U8 CTaBsicarpm, mature root
system.
B. AstragaiUB CTaBsicarpm, showing root of
young plant.
C. Baptisia bracteata.
PLA-TE 10.
A. Psoralea ttmuijlora, the tap root decayed.
B. P80ralea argophylla, showing entire root
in center.
PLATE 11.
A. BrauneM pallida.
B. Lygodesmia juneea, in two sections.
C. Lespedeza capitata.
PLATE 12.
A.. CeanothUB ovat'U8, root of a thirteen-year
old plant; a' is a continuation of a.
B. Amorpha canescens; a' is a continuation
of a.
PLATE 13.
A. Symphoricarpos w/garia, showing fine
network in surface soil.
B. Rhizomes and runners of Symphoricarpos
wlgaria; the horizontal line is the
ground line.
PLA-TE 14.
A. RhUB glabra, a portion of the root network
with ascending rootlet a.
B. Rh'U8 glabra, with ascending rootlets.
PLA-TE 15.
A. Coryl'U8 americana, the roots shown in
two sections.
B. Rosa arkamana, the roots shown in two
sections.
C. CorylUB americana, rhizomes and roots
PLA-TE 16. .
A. R08a arkansana, showing method of
propagation.
B. Rh'U8 glabra invading subclimax prairie.
PLATE 17.
A. Yucca glauea, showing the multicipital
stems and rhizome habit.
B. Prairie of southeastern Wasbington.
C. Meter quadrat in the prairie, showing
BaZsamorhiza, Festuea, LitMspermum,
and Hieracium.
PLATE 18.
v
A. The plains association near Colorado
Springs, showing Ariatida vurvurea
bunches in Bouteloua gracilis turf.
B. Psoraiea tenuijlora, in two sections.
C. Yucca glauea.
PLATE 19.
A. Stipa comata.
B. Petalo8temon candidus.
C. Eriogonum jamesii.
PLATE 20.
A. Short-grass plains, showing B01deloua
gracilis and Ovuntia polyacantha.
B. General view of the sandhill community.
VI LIST OF ILLUSTRATIONS.
PLATE 21.
A. A sandhill co=unity, showing Red-
jieldia, Petalostemon villo8us, Psuralea
lanceolata, and Chry80psi8 villosa, with
a socies of Eriogonum microthecum
at the left.
B. Roots and rhizomes of Calamovilfa
longifolia.
PLATE 22.
A. Andropogon hallii.
B. Muhlenbergia punuens.
PLATE 23.
A. Consocies of Aletes acaulis on the gravel-
slide, with Krynitzkia virgata in the
foreground.
B. Quadrat on the gravel-slide, showing
detail of surface.
PLATE 24.
A. Aletes aeaulis.
B. Network of fine rootlets of
C. Smilacina stellata.
D. Thlaspi alpestre.
PLATE 25.
A. Half-gravel-slide, showing Elymus trit-
icoides and the large bare intervals.
PLATE 25-continued.
B. Elymus triticoides.
C. Rubus delicio8U8.
PLATE 26.
A. Kreleria cristata.
B. Calamagrostis purpurasC6ns.
C. PotentiUa arguta glandulo8a.
PLATE 27.
A. Pieea engelmanni consociation, showing
the forest floor.
B. Quadrat in the same spruce forest, show-
ing Haplopappus parryi, Fragaria
virginiana, Thalictrum fendleri, etc.
PLATE 28.
A. Arctostaphylos uM-urBi, showing a por
tion of the root system.
B. Ribes lacu8lre, seven years old.
PLATE 29.
A. Root systems of ecads of Smilacina BIe1--
lata; a, gravel-slide; b, spruce forest.
B. Bouleloua gracilis from the sandhills.
C. Abronia fraorans from the plains.
PLATE 30.
A. Opulaster opulifolius, twelve years old.
B. Chrysopsis villosa from the sandhi1ls.
TEXT-FIGURES.
1. A meter quadrat in the Bouleloua com_PAGE 1
munity dominating a gravelly
ridge at Belmont, near Lincoln; B,
Bouteloua gracilis; A, Andropogon
fureatus; S, Sporoboluslongifolius;
M,Meriolixserrulata; Am, Amor-
pha canescens. . . . . . . . . . . . . . . . . . 9
2. Root system of Liatris punctata..... 10
3. Root system of Kuhnia glutinosa. . .. 10
4. Graphs showing the average daily
temperature (heavy line) and
humidity (light line) in the
prairie during 1916......... . . .. 22
5. Graphs showing the average daily
evaporation rates in the prairie
during 1916 (heavy line) and 1917
(light line) . . . . . . . . . . . . . . . . . . .. 22
6. Schematic bisect showing the root and
stem relations of important
prairie plants. This and figures
7 and 8 were drawn from photo-
graphs and data obtained by the
excavation and examination of
325 root systems of these 18
species: H, Hieracium 8couleri:
K, Kreleria cristata; B, Balsam-
orhiza sagittata; F, Festuca ovina
ingrata; G, Geranium viscosi8-
simum; .p, Poa sandbergii; Ho,
H oorebekia racemosa; Po, Poten-
tilla blaschkeana. . . . . . . . . . . . . .. 32
7. Schematic bisect: S, Sieversia ciliata;
W .. Wyethia amplexieaulis; LI,
Lupinus leucophyllus; Lo, Lupi-
nus ornatus; P, Poa Bandbergii;
E, Leptotrenia multijida; A,
A(lropyrum spieatum. . . . . . . . . .. 34
PAGE
Schematic bisect: Hu, Heuchera gla-
bella; A, Aslragalus arrectus; S,
Sidalcea oregana; H, Helianthella
douglasii; Ag, Agropyrum spica-
tum .......................... 36
9. Aristida purpurea.................. 47
10. Artemisiafrigida................... 47
11. Gutierrezia sarolhrre, showing wide
lateral spread of roots in the
surface soil. . . . . . . . . . . . . . . .. 48
12. A small part of caudex of Yucca (llauca,
showing number and extent of
horizontal roots, many exceeding
25 to 30 feet in length. . .. . . . .. 48
13. Argemone platyceras. . . . . . . . . . . . . .. 54
14. Lithospermum lineari/olium...... . ... 54
15. Lygodesmiajuncea........... ....... 56
16. Aragalluslambertii.......... . . . .. .. 56
17. Petaloslemon purpureus. . . . . . . . . . . . 58
18. Ratibida columnaris. . . . . . . . . . . . . . . . . 59
19. Senecio aureus oblanceolatu8. . . . . . . . .. 60
20. Underground parts of 1!erti-
ciliata pumila. . . . . . . . . . . . . . . . . 61
21. Opuntia eamanchiea, showing vertical
anchorage roots. . . . . . . . . . . . . .. 62
22. Top view of surface roots of Opuntia
eamanchiea, showing the wide
lateral spread. .. . .. .. .. . . .. ... 62
23. Red/ieldia j!exuosa, showing rhizomes
and root distribution. . . . . . . . . 69
24. Eriogonum microthecum . ..... '. . . . . . . 72
25. Root system of a tcn-year-old Arte-
mi3ia lili/olia . ........ ' . . . . . . .. 74
LIST OF ILLUSTRATIONS. VII
PAGE
26. Trode8oontia . . . . . . . . . . .. 74
27. Root system of Pelawslcl/urn 75
28. Gilia lcmoiJlora. . . . . . . . . . . . . . . . . . . .. 76
29. Underground parts of Psoralea lanew-
lala. showing root tubercles at a
depth of eight feet. . . . . . . . . . . . . 7i
30. 1 polllaa /eptophlllb. showing a portion
of the very extcn.i,'c root system 78
31. Krynilzkia The heavy slanting
line indicates the actual ground-
line.... ..... .. .. . . . . . . . . . . . .. 81
32. Paronychia jaflln;i... . . . . . .. . . .. . .. 82
33. Surface view of 1\ single root of Para-
nuchia jame8ii at a depth of 2 to
2.5 inehes. . . . . . . . . . . . . . . . . . . . . 83
34. Apocynum androsremijolium, showing
rhizomes and dense network of
roots ........... '.' '" .. . . .. ... 84
35. Pachywphua C<rspito8U8 ., . . . . . . . . . . .. 86
36. Eriooonum /lavum.... . .. . . . . .. . . . .. 87
37. Quadrat-bisect showing root relations
of gravel-slide plants. 'This was
made along the front of quadrat
shown in Plate 23, B: P, Paro-
n1lchia jame8ii; A, A/etes acaulis;
K, Krynitzkia virgata........ ... 90
38. Solidago oreop1iila, showing wide-
spreading lateral and deep cen-
tral roots. . . . . . . . . . . . . . . . . . . .. 93
PAGE
39. Bes8eya plantaoinea, showing the wide-
spreading root system. . . . . . . . .. 94
40. Root system of Geranium ccespitosum. 96
41. Root system of Frasera 8peeiosa . ... " 97
42. Pirola ehloranlha ..... .............. 100
43. Thalielrum lendleri ... .............. 101
44. Rhizome and roots of Erigeron asper.. 102
45. Fragaria virginia Ita . . . . . . . . . . . . 102
46. Allium cernuurn. . . . . . . . . . . . . . . . . . .. 102
47. Rhizomes and root system of Aralia
nudicaulis. . . . . . . . . . . . . . . . . . .. 103
48. Senecio cernuua. . . . . . . . . . . . . . . . . . .. 106
49. Heuchera parvi/olia ............... .. 107
50. Hapwpappua parryi. . . . . . . . . . . . . . .. 107
51. Root systems of eeads of Chamre-
nerium angustilolium: a, gravel-
slide; b, forest ................. 111
52. Fragments of the roots of Yucca
glauea: a, plains form; b, half-
gravel-slide form ............... 114
53. Allionia linearis from the plains. . . .. 115
54. Allionia lineaNs from the sandhilk .. 115
55 . 4bronia Iraorans from the sandhills.. 116
56. Chry80psis villosa, showing various
stages of development in the
plains soil.. . .. .. .. . . . . . . . . . . .. 118
57. Euphorbia montana from the plains ... 120
58. Euphorbia montana from the half-
gravel-slide. . . . . . . . . . . . . . . . . .. 120
THE ECOLOGICAL RELATIONS OF ROOTS.
INTRODUCTION.
Although considerable progress has been made in the study of root
systems of desert plants (Cannon, 1911, 1913; Markle, 1917), very
little information is available concerning the roots of other native
species. During 1914 the writer investigated the root systems of the
prairie plants of southeastern Washington, where the annual pre-
cipitation is only 21.6 inches and occurs mostly in the period of rest:
Since that time it has been planned to make a comparative study of
the roots of prairie plants growing in a more humid region and where
the precipitation occurs mostly during the season of plant growth.
The opportunity for such study came during the fall of 1917 and work
was carried on vigorously until the soil b ~ c a m e frozen in December.
The study was resumed early in the following spring. In June 1918
the field of investigation was extended to the Great Plains and sand-
hill region of Coloraoo, while later in the summer a large number of
plants were excavated and examined in various habitats about Pike's
Peak in the Rocky Mountains.
A knowledge of root distribution and root competition under dif-
ferent natural conditions is not only of much scientific value, but it
also finds practical application in a better understanding of the value
of plants as indicators for distinguishing lands of grazing value only
from those with possibilities of crop production. It will result in a
more intelligent solution of the ecological probleins of grazing and will
likewise be of great aid to the forester in selecting sites for afforesta-
tion. Moreover, a knowledge of root distribution will throw a flood
of light upon many of. the problems of plant succession. Indeed, the
phenomena of ecesis, competition, and reaction can not be completely,
if indeed correctly, interpreted without a knowledge of the extent,
position, and relation of the root systems of the plants.
Shantz has given us an excellent example of the value of a knowl-
edge of root distribution in his study of the natural vegetation of the
Great Plains as an indicator of the capabilities of land for crop produc-
tion (1911). Sampson (1914, 1917) has made a study of root systems
of many range plants in considering their life history, forage value,
and the natural revegetation of range lands; while foresters are just
beginning to study the roots of plants as indicators of conditions of
soil moisture on various sites (Korstian, 1917).
This paper contains descriptions of the character, depth, and dis-
tribution of the roots of about 140 species of plants. These include
shrubs, grasses, and other herbs, a few of the latter being noxious
1
2 THE ECOLOGICAL RELATIONS OF ROOTS.
weeds. It represents the results of the examination of approximately
1,150 individual plants in 8 different communities, as follows: prairies
of eastern Nebraska, chaparral of southeastern Nebraska, prairies of
southeastern Washington and adjacent Idaho, plains and sandhills of
Colorado, the gravel-slide, the half-gravel-slide, and forest communi-
ties of the Rocky Mountains of Colorado.
The method employed in excavating root systems was to dig trenches
2 to 3 feet wide and 6 to 10 feet long to a depth of about 6 feet,by the
side of the plants to be examined. This offered an open face into which
one might dig with a hand pick furnished with a cutting edge on one
end, and, after sufficient practice and acquaintance with the soil
texture, successfully excavate a root system almost in its entirety.
Of course, the trenches were deepened as work progressed and the
working level sometimes reached a depth of 10 or even 16 feet (plate
1). A total of more than 100 such pits was used. To assure absolute
certainty as to maximum depth, for extreme care must be used in
excavating root termini, the soil underlying the deepest roots was
. usually undercut about 12 to 18 inches below the root-ends and care-
fully examined as it was removed. For the deepest-rooted species of
herbs and shrubs it was found expedient to have a longer trench with
two levels, one at about 7 to 9 feet and a second one about twice as
deep. Thus the soil could be removed from the lower to the higher
level as work proceeded. Indeed, in several cases where the roots
extended to depths of 18 or 20 feet or more (for example Rosa arkan-
sana or Lygodesmia juncea), the deeper soil was removed by means of
a bucket attached to a rope. Considerable danger from caving was
experienced, especially in the sandhill soils and also in the deeper loose
loess soils as well. In fact, it was found inexpedient to remove the
entire root system in a few cases. In each community the work
extended over a field sufficiently large, sometimes several square
miles in extent, so that any local differences in soil texture, etc., were
eliminated.
All of the roots examined, except as otherwise indicated, were of
mature perennial plants. The practice followed was to examine
several roots of a given species and then to write a working descrip-
tion of the root system. These descriptions were kept at hand, and as
new roots of the same species were studied, any variation from the
original description was carefully noted. While many of the root
systems, especially those of the grasses, were removed in their entirety
and photographed against an appropriate background, and a few
photographed in position, many others were drawn in place. The
sketching was first done with pencil on a large drawing-sheet ruled
to scale. Drawings were made simultaneously with the excavating of
the root and always to exact measurements. When entirely com-
pleted they were retraced with India ink. Such a drawing often repre-
INTRODUCTION. 3
sents, the extent, position, and minute branching of the root system
more accurately than a photograph. This is especially true of the more
extensive root systems, for here, even under the most favorable con-
ditions, the photograph is always made at the expense of detail, many
of the finer branches and root-endings being obscured.
During the course of these investigations the writer has become
greatly indebted to the following persons for faithful assistance in the
pursuance of this work. It is a pleasure to acknowledge here the
indebtedness to my students, Messrs. F. C. Jean and Alvin Goke, for
much help in the excavation of the plants, and to Miss Annie Mogen-
sen and Mrs. F. C. Jean for their assistance in drawing many of the
root systems. I wish also to acknowledge the helpful suggestions and
encouragement given by Dr. F. E. Clements and Dr. R. J. Pool
throughout the period of the work. To Professor T. J. Fitzpatrick
I am also indebted for careful reading of the and proof.
I. THE PRAIRIES}
The prairies of eastern Nebraska in which. these studies- were made
are too well known to need discussion here (plate 2). A general floristic
description has been given by Pound and Clements (1898, 1900), a
more detailed one for a portion of southeastern Nebraska by Thornber
(1901), and recently a more specific account of the local region under
consideration by Weaver and Theil (1917). As the various species are
considered, brief comment will be made upon their relative importance.
Therefore, we may proceed to a consideration of the rQot habits of the
individual species and then examine the environmental conditions
under which the plants grow. Finally, the correlations between root
habit and environment will be discussed. This sequence will be fol-
lowed for each plant community. .
Panicum virgatum.-Although showing a preference for loose sandy soils,
panic grass grows abundantly in many situations throughout the prairies.
Its size, abundance, and duration combine to make it an important com-
ponent of the prairie vegetation, where it often holds the rank of a dominant
species ..
This grass has the longest root system of any species examined. The roots
are very coarse, many having a diameter of 3 or 4 mm. They pursue a verti-
cally downward course, spreading only a little near the surface, to a maximum
depth of over 9 feet. Several plants reached depths of over 8.5 feet (plate
4, A). In the first 6 or 7 feet of soil the r ~ t s are very little branched, the
laterals (usually less than 3 inches long) occurring only scatteringly. In the
deeper soil numerous fine laterals occur, although the last 6 to 12 inches are
often very poorly branched and the roots end abruptly. The main roots
remain nearly unifonn in diameter for most of their course or taper so slowly
that at 5 or 6 feet they may still have a diameter of 2 mm. The type of soil
considerably modifies not only the root penetration but also the manner of
branching. The plants that were growing in a gravelly soil with a sandy
subsoil underlaid with an impervious blue clay were found to penetrate only
7 feet to the clay, but the branching was much more pronounced. The cortex
on younger roots is pearly white and very brittle; on older ones smooth,
pinkish white, and of a papery consistency. Eight plants were examined.
Andropogon furcatus.-Tbe two bluestems are dominants among prairie
grasses. They fonn a large part of the prairie hay of Kansas and Nebraska.
The taller and deeper but coarser-rooted A. furcat'U8 is less resistant to drought
than the shorter, finer-rooted A. scopari'U8, as is shown both by their local
and general distribution. Throughout the prairies the forfner luxuriates in
the draws and on lower lands, while the latter dominates higher areas.( West-
ward the big bluestem seen drops out, while little bluestem not only forms a
. ""dominant of widest distribution and most controlling influence in the bunch-
. grass association of the sandhills- (Ppal, ] 914;..224).. and If enters into dis-
turbed areas of the wiregrass association in ColoradO"" (Shantz, 1911: 52),
but also occurs on rough hillsides where water penetrates readily, throughout
much of the area included in Wyoming and Montana. .
1 The major portion of the work in this section was done by the writer in conjunction with Mr.
Lyman H. Andrews, who voluntarily joined the colora in December 1917, and to whom joint
credit ill herewith acknowledged.
THE PRAIRIES.
5
Ttm Eyck (1904: 216) examined the roots of A. furCldus at Manh!l'ttan,
nsas and found that they "form a dense, tough sod, from 6 to 8 mches
ck the subsoil is filled with a great mass of roots." "The roots were
off at 4.5 feet from the surface, but from their size they must have
extended at least 2 feet deeper into the compact clay subsoil."
J'welve plants were examined near Lincoln. The very roots
both vertically and obliquely downward, a few almost honzontally,
an once thoroughly occupy the soil and fonn a dense sod. The roots(
extend obliquely away from the bunches to more than a foot before turnmg
wnward. The larger roots vary from 0.5 to 3 mm. in diame!-er and may
ch a depth of 6 feet and 10 inches. Most of the plants exammed reached
&epths of over 6 feet {plate 3, A). In locations where-a-hard clay subsoil
occurred the roots were 21io 2.6 All of the roots branch prof?SC1y,
main laterals being from 2 to 6 mches long. However, here agam
IOunt of branching and the length of the laterals are closely correlated
U soil texture, always being less in baM soils. The roots taper so gradually
that at 4 feet thay are nearly as large as at the surface.
j
Pied to a depth of 5.feet. The ends of the roots are extremely well
ched to the very tip. In color the roots are reddish-brown. They have
ery loose papery cortex which is easily removed and reveals the tough,
yellowish stele,
seoparius.-This grass is figured by Shantz (1911: 56) as

ding to a depth of about 5 feet and thoroughly occupying the sandy soil
the bunchgrass association of eastern Colorado. It is interesting to note
that the deep-rooted Panicum virgatum also occurs here.
In these studies plants of this species were examined in two soil types. The
lit group was in porous, gravelly soil mixed with sand and underlaid with a
tobky subsoil of decayed sandstone at a depth of 3 feet. The other group
grew in clay-loam soil with a clay subsoil. In the fonner habitat none of the
'lts of the several plants examined reached depths greater than 28 inches,
!lile in the clay loam several plants had a maximum root depth of about 65
wehes (plate 3, B). The roots are much finer than those of A. furcatus, being
only 0.1 to 0.8 mm. in diameter. The lateral spread of the roots is very
. milar but somewhat more pronounced. Roots are so abundant as to fonn
. dense sod, completely filling the soil to a depth of from 12 inches in gravelly
Uil and to as much as 30 inches in clay loam. The surface is especially well
occupied with dense masses of finely branched rootlets. All the roots branch
IIWI'Ofusely to the third or fourth order, many of. the branches being over 30
ches long. The deeper soil (from 30 to 60 inches) is fairly well occupied .
...Ale roots are light-brown in color and have a very thick cortex which peels
f!l.. thus exposing the tough yellowish stele. The deeper roots are
..,....ter ill color,1>rofusely branched, and very brittle.
Andropogon nutans.-Goldstem is a dominant in the subclimax prairie.
"Oke the othe! andropogons and Panicum virgatum, it also matures late in
It 18 one of the deeper-rooted prairie grasses. Of the 5 plants
l:amined the maximum depth of root varied between 51 and 59 inches
3, c). These were growing in a clay-loam soil which extended to a
2 pth of 3.5 feet and below which occurred pure sand. The roots vary from
mm. to less. than 0.5 mm. in diameter. They are very abundant, spread
\terally but and occupy the soil, branching profusely to the
Klond and third order. WIthin a foot from the surface, however, most of
.....e roots become less than 0.5 mm. in diameter, forming a dense network to
a depth of about 3 feet. Even in the fourth foot the roots are quite numerous,
many of them breaking up at this depth into clusters of small branches. The
roots are slightly reddish in color. The branching is characteristic but difficult
to describe. .
Stipa spartea.-This perennial grass is one of the dominants in the prairies,
being especially conspicuous during the month of June, when it gives tone to
the estival aspect. Its root system is rather meager when compared with
most of the other prairie grasses. Strong fibrous roots from 1 to 1.5 mm. in
diameter descend rather vertically into the soil to a maximum depth of only
21 to 26 inches (plate 4, B). A few spread laterally in a diagonal direction to
a distance as great as 10 or 12 inches. Numerous smaller roots fill the surface
soil, while the larger ones send off many laterals to a depth of about 14 inches,
where the main roots may break up into many fine branches. The hairy
cortex causes the soil to cling to it very closely. This pulpy cortex has a
grayish-white color and peels off very easily, leaving a very wiry, tough stele.
Rooleria cristata.-Whether onthe prairies of Minnesota or Nebraska or
in the Pacific Northwest, Kreleria is an important grass in the estival aspect.
As in the case of Stipa spartea, which also makes a rapid growth and blossoms
early, the vegetative and reproductive activity may be correlated with the
shallow root system. In eastern Washington, where Kreleria is a common
bunchgrass, it flowers in late June, soon dries up, and remains dormant until
revived by the autumn rains. In this region the deepest root found was at
28 inches, and 15 inches was determined as the average depth (Weaver, 1915).
This plant has a very shallow but exceedingly well-developed root system.
None of the roots of the 7 plants examined reached depths of over 21 inches,
while the average root depth was about 15 inches (plate 5, A). A great abun-
dance of fine rootlets spread out from the base of the plant and occupy the
soil exclusively for a distance of 8 inches on each side of it. Indeed, some of
the roots run almost horizontally and are less than 0.5 inch deep in the soil.
The roots are yellowish-brown in color, usually less than 0.2 mm. in diameter,
and branch and rebranch to form a dense mat.
Elymus canadensis.-This plant is an important prairie species of wide
distribution. Like Kreleria, it is comparatively shallow-rooted, but has a
widely spreading root system. Five plants in loess soil gave maximum root
depths of 16, 17,20,21, and 22 inches respectively. Plate 5, B, shows the
rather meager root development when compared with most of the other
grasses. The obliquely running roots reach distances of 20 inches or more
on either side of the base of the plant. Lateral rootlets are seldom over 2
inches long and may branch to the third or fourth order. The roots are white
in color and from 0.1 to 0.5 mm. in diameter. They are very tough and wiry.
Agropyrum repens.-From the extensive rhizomes of this perennial, which
are often 2 to 3 feet long, arise numerous fine, silvery-white roots. These
send off rather poorly branched laterals as they descend somewhat vertically
downward, some to a maximum depth of 8 feet. The first 30 inches of soil
is abundantly filled with roots, while many penetrate to a distance of 4 to 7
feet. Many of the roots of the 10 plants examined pursued a peculiar zigzag
course, a character which was more pronounced in the deeper roots.
Distichlis spicata.-This low, direcious perennial of seacoasts and alkaline
soils is still quite abundant on the "salt flats" about Lincoln, although it is
being replaced by other species as the soil becomes less salty, due to better
drainage resulting from the straightening of stream courses. It has consider-
able forage value throughout the West, where it occurs abundantly in low
saline situations. About Lincoln most of the area occupied by salt grass is
THE PRAIRIES.
7
alluvial wash, the various soil strata. showing quite (plat;e
1, B). The plants examined were growmg m pure A chenncal
nation of the first 8 inches of soil (the samples being taken soon after a
had wet the soil to just this depth) gave salt of 2:6 per cent.
is less than in the adjacent and succeSSIonally earlier Atriprex zone, which
gave 3.1 per cent alkalinity. In both cases the greater part of the salt was
Sodium carbonate.
1
h' . f 1
In the 8Uccession it is repla.ced by Agropyrum repen8, whic m turn IS 01-
lowed by SporobolUB longifoli'lJ.!, Panif:um uirgat!,-m, Andropogon furcatUB,
and other prairie plants. An mterestmg SUCceSSIOnal sequence was deter-
mined in a rich alluvial flood-plain, where the water-level occurred in gumbo
soil at a depth of about 6 feet. Although only an occasional specimen of
DiIlichlia was to be seen, the soil contained three distinct strata of the abun-
dant and well-preserved rhizomes at depths of 13, 9, and 6 inches respectively.
These indicated successive overflows and depositS. Above these the soil was
filled with a dense network of the rhizomes of Agropyrum repen8. However,
only a few of these plants were still alive, the soil being almost completely
occupied by alternate areas of Sporobolm longifolim and Bulbilis dactyloide8.
Diatichlis has better developed rhizomes than almost any other grass
examined. They vary from 2 to 5 mm. in diameter, often being somewhat
flattened parallel with the soil surface. One specimen was found to be 9 feet
long. It supported 19 tufts of plants. The rhizomes are exceedingly tough,
being covered with a hard, shell-like cortex. The terminal buds are long and
sharp-pointed, as are also the leaf-scales at the regularly spaced nodes which
are about 2 inches apart. The rhizome depth is quite uniform at from 4 to 6
inches. The rhizomes branch extensively, sending off laterals in all directions.
The roots are relatively shallow, but few occur in the first 4 inches of soil.
They branch from the rhizomes in both vertical and horizontal directions,
and while the lateral spread is not great, they occupy the soil thoroughly to .
a depth of about 18 inches. Relatively few reach a depth of over 2 feet.
They are from 1 to 3 mm. in diameter and branch rather sparingly to the
third order, these laterals being only 1 or 2 inches long. The deeper roots are
even more sparingly furnished with laterals, which are only 1 to 3 mm. in
length. The yellowish-white cortex of the older roots is thick and pulpy.
When removed it reveals a pearly-white stele. Thirteen plants were examined.
Sporobolus longi!olios.-This perennial, late-maturing grass is often an
important component of prairie vegetation, especially in the earlier stages of
development. Although it occurs in typical prairie, it is often more abundant
on eroding banks along roadsides and in disturbed areas generally from dry
hill-tops to alluvial flood-plains. The short, thick rootstocks spread in all
directions and form bunches varying from 4 to 18 inches in diameter.
The rather coarse, fibrous roots penetrate the soil to an average maximum
depth of only 24 inches. The greatest root depth of the 17 plants examined
ranged from 17 to 40 inches. However, the roots are very dense and
occupy all of the soil, spreading laterally from the base of the plant
In an abnost horizontal direction to a distance of from 12 to 20 inches (plate
6, A). Thus an area of soil of 6 or 7 square feet may be thoroughly occupied
to a depth of 18 inches by the roots of a single bunch. The roots vary from
1 to 2 mm. or less in diameter, many of them keeping the original diameter
to a depth of 18 inches. They are pearl-white in color, very tough and wiry,
and the cortex is densely covered with short hairs. When the cortex is
removed it reveals the thick, white stele. All of the roots branch profusely,
I The writer is indebted to Professor C. J. Frankforter for these determinations.
but the laterals are only 0.5 to 2 inches long. These often branch again. The
roots below the IS-inch level are relatively few and the soil is not well occupied.
Roots were examined in 3 different soil types and found to be very similar in all.
Aristida oligantha.-This species is abundant on prairies, especially in drier
soils. It frequently dominates areas where the soil is shallow, or plays the
r ~ l e of an interstitial among the taller grasses. The root systems of 10 plants
of this annual grass were examined. They were growing in a clay-loam soil.
The white, threadlike roots are less than 0.2 rom. in diameter and are found
abundantly only in the first 10 inches of soil. While some of the longest
reach depths of 40 inches, relatively few occur below 18 inches. They are
so fine and brittle that it proved impossible to secure material for a photo-
graph. Compared with other grasses the roots are much less abundant.
Bulbilis daetyloides.-Perhaps no grass of the western plains, except
grama, is better known for its valuable characteristics as a pasture grass than
the buffalo grass. In the region of this study it can not compete with the
taller prairie plants. However, it is found in dense patches resulting from its
method of propagating by stolons, on low-lying lands where water may stand
for a time in "pockets" or where over-grazing has killed the taller grasses.
Ten Eyck (1904), working at Manhattan, Kansas, and in a soil of which the
surface foot was a mellow, dark loam, underlaid with a rather compact, clayey
subsoil, found "the roots are numerous but they do not penetrate deeply
into the soil". Shantz (1911: 38), dealing with the hard soils of the Great
Plains region of Colorado, where "the soil is only rarely wet down to a foot
or more," states that "almost the entire root system of the short grasses
(Bulbili8 dactyloide8 and BtlUteloua oligo8tachya) is limited to the first 18 inches
of soil. They have a very extensive surface root system." Robbins (1917:
70) states that roots of buffalo grass sometimes go to a depth of 7 feet. This
latter statement checks up more nearly with our findings. '
A dozen root systems of this grass were examined in two different areas,
both of which were in alluvial soil on bottom lands. The roots are very fine,
the largest being less than 0.5 rom. in diameter. They scarcely spread at all
laterally, but form a dense mat to a depth of 12 to 18 inches, branching mostly
only to the second order. These hairlike laterals are usually not over 0.5
inch in length. Although these roots are very fine, they are quite tough and
easy to follow. At 18 inches depth they become less numerous and are poorly
branched, so that the deeper soils are sparsely occupied. However, many
roots occurred at 4.5 feet; and numerous others continued vertically downward
to a maximum depth of from 58 to 73 inches and in one of the trenches in
gumbo soil to the ground-water level. The shallower roots are light-brown
in color, while the deeper hairlike roots are nearly white. _.
Bouteloua graeilis.---Grama grass occurs often in fairly pure stands on the
lighter soils of gravelly ridges and it is also frequently found dominating
areas of alluvial soil on bottom lands. Figure 1 shows a typical area domi-
nated by Bouteloua into which Andropogon furcatm is invading. The soil is a
very porous coarse sandy to gravelly loam deposited by glacial action, with a
water-holding capacity of about 40 per cent of its dry weight. This grama
consocies, in which a number of root systems were examined, dominates a
ridge at Belmont near Lincoln, which is entirely surrounded by tall prairie
grasses rooted in clay loam. The transition from one soil type to the other is
very abrupt and the ecotone is correspondingly sharp. Frequent determina-
tions invariably showed a higher available water-content in the clay-loam soil.
The root system is extremely well developed, great masses of fine roots
occupying every cubic centimeter of soil to a depth of 18 inches. A few roots
THE PRAIRIES.
9
reached a maximum depth of 46 inches, although below 2 !eet. the roots are
very sparse (plate 6, B). The largest roots are only 0.5 mm. ill dia.D!-eter. The
laterals are usually not over an inch long. These abundant threadlike laterals
branch only poorly. The roots are brownish white in col?r and of a fir?1
texture. Other groups of plants examined in two locations on alluvial soils
showed a somewhat poorer development of the root system, but the
distribution and depth were very similar to those growing in the gravelly soil.
This root distribution stands quite in .
contrnst to that found by Shantz !n Col-
orado. This would seem to be a clear
case of environmental conditions pro-
foundly modifying root development.
Llatris the numerous
blnzing-stara which add beauty to the
prairies in autumn, L. punctata and L.
soorioso are probably the most abundant.
The xerophytic nature of the former and
its extensive range far into the drier
westward may be explained in
part by its deep root system.
Seven individuals of this species were
excavated. All had strong tap-roots.
Those growing in clay with a sandy sub-
soil reached depths of 11 feet 5 inches FIG.1.-A meter quadratintheBmde-
and 15 feet 9 inches respectively, while loua community dominating a
three growing in clay-loam with a clav gravelly ridge at Belmont,
J near Lincoln.
subsoil penetrated only to a depth of
about 6 feet 8 inches. A diagram of one B, Boutelou.a gracili8.
of the larger specimens (fig. 2), recon- A, Andropogonj'Urcatus.
structed from a photograph and from a S;r::;us =iUS.
made to scale in. the. field,
pictures the roots as occurrmg ill one ' .
plane. On other specimens small laterals were very scarce to a depth of 9
feet, while at 12 feet the soil was filled with small, silvery white, sparsely
branched rootlets. This scarcity of absorbing laterals was again noted in the
clay-loam soil of the high prairie until a depth of about 6 feet was reached.
Even here the branching was not so pronounced as on the specimens in the
lighter soil. The older roots are woody and chocolate-brown in color.
Liatris scariosa.-This plant, unlike L. punctata with its strong deep tap-
root, is characterized by a large woody corm, 3 to 5 inches in diameter, from
which arise very numerous fibrous roots. These are only 1 to 2 mIn. in diam-
eter, but they are very finely branched and spread laterally, thus thoroughly
oC,cupying the first 2 feet of soil. The depth of penetration was not deter-
nuned.
Solidago rigida.-This goldenrod is not only abundant in the less xero-
ph,ytic prairies of Minnesota and Nebraska, but extends far westward into the
drIer ,grasslands, In Wyoming and Montana its presence, together with
other species" invariably indicates rather permeable soils with at least
a frur amount of available water.1 .
1 During the summer of 1917 the writer, while working on grazing problems in the prairies of
the .Northwest, had an excellent opportunity to trace the westward extension of many species
of the less xerophytic prairies of eastern Nebraska and Minnesota far into the Great
Plam8: Indeed, of the two associations of the prairie-plains climax of Clements (1916),
tBhe Bhpo-AI/TOPl/TUm prairie occupies much more territory in the Northwest than the Bullnlill-
outeloua plains,
10
This plant has a root system which spreads immediately below the surface.
Many of the numerous small roots pursue an oblique direction and spread
from 12 to 18 inches on either side of the plant before they turn downward.
Most of the roots are only about 1 mm. in diameter and poorly branched .
f- 1'-
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Clay
Sandy
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__ __
FIG. 2.-Root system of Liatria punctala. FIG. 3.-Root system of Kuhnia gluti1W8a.
However, they are very abundant in the first 2 feet of soil, while maximum
depths of over 5 feet are attained. They are yellowish-brown in color and have
a smooth cortex, except the lower roots, which are silvery white. Only 4
plants were examined.. .
THE PRAIRIES. 11
Solidago canadensis.-This abundant prairie goldenrod propagates by
means of strong woody rhizomes, a centimeter or less in diameter, which run
horizontally about 2 inches below the soil surface. A13 in S. rigida, the roots
are fibrous. They are about 1 to 2 mm. in diameter and pale yellow in color.
They descend rather vertically into the soil. From these abundant roots
numerous laterals are sent off, beginning just below the surface and con-
tinuing to a depth of from 9 to 10 feet. However, the main roots keep their
identity throughout their course and are very poorly branched throughout
the last foot. Thus the soil was well occupied with the roots of this species
and few other roots were found in the Solidago area. Eleven plants were
examined, most of which gave a maximum root depth of 10.5 feet, while one
reached 11 feet.
Silphium laciniatum.-The wide distribution of this coarse perennial
prairie plant is well known. It has a large, fleshy, deep tap-root. The root
diameter just below the multicipital crown, which gives off numerous shoots,
varied from 1 to 2 inches in the 5 plants examined. The tap-root descends
vertically and tapers so rapidly that at 3 feet it may be only 0.5 inch in
diameter (plate 7, A). It pursues this general vertical direction to a maximum
depth of from 9 feet to 13 feet 8 inches, the last 2 or 3 feet of its course being
somewhat tortuous. The whole root is a dark reddish-brown in color, the
cortex of the first 18 inches being distinctly ridged and wrinkled. Scattered
laterals, 2 mm. or less in diameter and from 6 to 18 inches long, are present
on this upper portion of the root. There are relatively few large laterals.
These frequently run off horizontally for a distance of 3 or 4 feet and then
turn downward. The deeper roots are also fleshy and brittle and end rather
abruptly; usually they are freely branched.
Amorpha canescens.-This half-shrub is a characteristic subdominant of
the summer aspect on the prairies of eastern Nebraska. Late in June it
begins to blossom and throughout July its leaden color gives tone to the
landscape. This perennial legume has a very extensive, woody root system
which reaches depths of 6.5 to 7.5 feet. While some of the plants examined
branched into large lateral roots just below the enlarged knotty crown, others
sent vertically downward a single tap-root. These extremely tough woody
roots have a maximum diameter of more than a centimeter. Frequently,
large laterals run off horizontally at a depth of only 4 to 6 inches below the
soil surface to a distance of 2.5 to 3 feet before they begin their downward
course (plate 7, B). Few laterals are given off until the roots reach a depth of
from 2 to 3 feet; then the roots by dividing become much finer and branch
quite profusely. The terminal branches are very fine, almost threadlike.
The roots are cinnamon-brown in color; the cortex is smooth, except on older
portions, where it is finely scaly. Five plants were examined.
Helianthus rigidus.-This sunflower, common on Nebraska prairies,
propagates by strong rhizomes, from which arise abundant fibrous roots only
. 1 to 1.5 mm. in diameter. These reach a depth of about 50 inches.
Brauneria pallida.-0nly 2 plants were exaniined. Both had strong,
vertically descending tap-roots which reached depths of 51 and 66 inches
respectively. These roots were 1 or 2 cm. in diameter. They were poorly
supplied with branches, only 3 or 4 short, weak laterals occurring on each root.
Even the tip is unbranched. They are woody, and are chocolate-brown in color.
Petalostemon candidus.-Both this plant and Brauneria are common and
often very abundant prairie species. Like Brauneria, it has a strong, deep,
rather poorly branched tap-root. Of the two specimens examined the smaller
I
had a root 7 mm. in diameter and a total length of 42 inches. The larger was
10 mm. in diameter and 68 inches deep. The laterals that do occur may
originate near the surface and are branched to the third or fourth order. One
plant sent off two strong laterals at a depth of 12 inches. The deeper roots
are very fine and quite well branched. All have a characteristic yellow color
throughout. .
Vernonia baldwinii.-This weed propagates by strong rhizomes, a centi-
meter or more in diameter, which occur about 4 inches below the soil surface.
The roots, which are very abundant (20 or 30 arising from a single plant), L
are very different from any other roots examined. They are tough, smooth,
yellowish-white, and unbranched throughout the first few feet of their course.
While the stele is only about 1 mm. in diameter, the fleshy cortex is thick
enoUgh to give a total root diameter of from 3 to 9 mm. Many spread to a
distance of 20 inches from the base of the plant before turning downward.
Thus in a weedy pasture the surface soil is filled with the shiny, white,
unbranched, rope-like roots. At about 3 feet in depth they begin to branch
and the branches become more numerous in the deeper soils. Most of the
finer branches and terminal rootlets, however, occur at 9 or 10 feet. Seven
plants were examined, all of which penetrated to a depth of about 11 feet.
The maximum depth recorded was 11 feet 6 inches.
Kubnia glutinosa.-This prairie plant "has deeper roots than most other
species examined. The maximum depths of 4 plants were 16 feet 8 inches, 16
feet 6 inches, 16 feet 10 inches, and 17 feet 3 -inches, respectively. A part of
one of these is shown in plate 8, A, where only about a third of the root
system is exposed. The root in figure 3 was reconstructed from the photo-
graph of another root and a diagram made to scale in the field while excavating
the plant. This diagram shows all of the larger roots, the extent of their
lateral spread, degree of branching, etc. Although single water-content
determinations without wilting coefficients are usually of little value, yet here
the wetness and dryness of the various soil strata and the corresponding root
branching were so marked as to warrant including duplicate determinations
made on November 5, 1917, after several weeks of very dry weather. The
clay layers at 8 and 12 feet respectively are more or less impervious, and
collect and hold water readily. The tap-roots vary from 1.2 to 3.5 em. in
diameter. All the roots are nearly white in color.
Verbena stricta.-This weed, very common in old pastures, has a strong
tap-root from 1 to 2 cm. in diameter, which reaches depths ranging from 3.5
to over 4 feet. Seven plants were examined. In all cases profuse branching
occurred from the soil surface to the extreme root-tip. Because of the numer-
ous strong laterals sent off from the tap at all levels, the latter decreases in
size rapidly. The&J& lateral branches lay hold of the soil to a distance of 18
inches on all sides of the plant. Small rootlets are abundant, the. whole
forming an extremely well-developed root system.
Grindelia squarrosa.-This weed has a strong tap-root abundantly supplied
with well-developed laterals. The main root varies from 0.5 to 1 inch in
diameter, but it gives off laterals so abundantly that this size is not long
maintained. Maximum depths of 50, 52, 40, and 73 inches were recorded for'
the 4 plants examined. There are many very fine roots, 3 to 9 inches long,
occupying the surface soil. The larger laterals begin to branch off in the first
foot of soil, and run off obliquely from the main root to a distance of about
2 feet. Like the tap, these are supplied with abundant rootlets, the whole
forming a very extensive absorbing surface. The roots have a light yellow color.
THE PRAIRIES.
13
Shantz list.8 Grindelia as one of the plants found in short-
grass association of Colorado. It usually indicates disturbed. conditIOns and
is especially abundant during wet years .. appears that root system
must be greatly modified under these conditions where the SOillS only rarely
wet below a foot or two. Indeed, it seems to thrive so well and under such
varied conditions that it is a conspicuous throyghout many of the
Western States. It is likely that the great plastlclty of lts root system makes
this wide distribution possible. .
Olycyrrhiza lepidota.-This characteristic legume has
rhiaomes eeveral feet long and a deep, fleshy tap-root. In the several SpeCi-
mens examined the tap-root varied from 0.5 to 2 inches diameter. From
the multicipital crown as many as 15 to 20 stems may anse. Just below the
surface fOil emalliateralfl, a few millimeters to a centimeter in diameter, are
often given off, in a more or less horizontal dire<)tion. These often .tum down
abruptly. At a depth of 1 to 3 feet the tap frequently breaks mto many
strong branches with a lateral spread of 2 or 3 feet from the base of the crown.
They run off rather obliquely from the main root (plate 8, B) giving rise to
numerous long branches. These tertiary roots are usually only 2 to 5 mm. in
diameter and rebranch poorly, a lack of small, absorbing roots being quite
characteristic. The deeper soils (8 to 10 feet) are thus well filled with small,
vertically descending, poorly branched roots, 1 to 3 mm. in diameter. Like
the older roots, these are of a light-brown color and fleshy but quite firm.
They shrink rapidly upon exposure to the air. At 10 to 12 feet, where these
rootlets entered the jointed clay-loess soil, they branched rather profusely.
The roots of several plants were traced to depths of 11 or 12 feet. One of the
larger ones was still 2 mm. in diameter at this depth and probably penetrated
several feet deeper. The odor is very characteristic.
Astragalus erassicarpus.-During May 1918, a large trench was dug ona
west hillside at Belmont, about 45 feet from a wet Spartina cynosuroides zone
in the valley. The surface foot of good loam soil gave way to about 3 feet of
exceedingly hard joint clay, in which roots (and especially those of grasses)
frequently followed the joints and branched largely (locally) in one plane.
The deeper soil became wetter and also somewhat sandy, but very gummy
and sticky. The water table was reached at about 7 feet. From this trench
Aatragalua, two species of P8oralea, and Baptisia were excavated.
ABtragalua crassicarpus forms extensive societies in the early spring. A
o! 3 ounger was first examined; the plants were very
similar m Slze, root diameter, branching, and depth. One (shown in plate
9, B). had a pronounced tap-root with a diameter of 1 cm. and ran almost
vertically downward to a depth of 6 feet, giving off a number of horizontal
branches, 1 to 2 mm. in diameter, near the surface, and extending laterally
only a few inches before turning downward. Other smaller laterals from 0.5
mm. to those hairlike in diameter occurred sparingly and at intervals of about
1 em. or less to a depth of 4 feet. Most of these branches were less than a
oenbeeamtimeter in length. Below 4 feet, in the softer, wetter soil, the branches
b -: m?re pronounced, being densely covered with root-hairs and often
ranching m two planes. However, none of these branches exceeded an inch
d The tip of the tap (as well as the tips of the laterals, which reached
epths only about half as great as the former) was no better branched than
the last 2 or 3 feet of the root. There is a very noticeable tendency for
e roots to follow the crevices in the joints of the clay.
An plant had a strong tap-root with a diameter of 1 cm. At a depth
of 2 em. It gave off a strong lateral 5 mm. in diameter and at about the same
I
level 3 other branches of equal or slightly smaller diameter (plate 9, A). They
ran off obliquely in 4 different directions, but none to a distance greater than
10 inches before they turned rather vertically downward; two more branches,
each 4 mm. in diameter, were given off at depths of 6 and 8 inches respectively.
All of these branches were very poorly supplied with laterals, as has already
been noted for the younger plants. They taper uniformly, so that at a depth
of 2 feet none had a diameter of more than 3 mm. The tap as well as several
of the ultimate branches of the laterals reached a depth of 6.5 feet. The
hairlike termini are sometimes unbranched, often for several inches, but are /
more often well-branched. Nodules 1 mm. or less in diameter occur at all .
depths, even near the root-tips. The roots are very brittle and hard to
recover from the joint clay. They are yellowish brown in color except the
older portions, which are dark brown._
Psoralea tenuiflora.-This very abundant legume forms societiel' over large
areas of prairie during June. Indeed, for a period these tall, coarse herbs quite
overtop the grasses. The one shown in plate 10, A, had a tap-root 3 cm. in
diameter, which ran vertically downward. At a depth of 27 inches it appeared
dead, but it was traced to a depth of 52 inches, where it was still 3 rnm. in
diameter. At a depth of from 1 to 2.5 feet many laterals occur. These are
from 7 to 12 rnrn. or less in diameter and frequently run off obliquely (as
shown in the figure) for distances of 8 to 18 inches before turning, often
abruptly, vertically downward. Several of these laterals, including some of
the flmaller ones, reached depths of 5 or 6 feet. The lateral spread is such
that at the tips many of these roots are at a horizontal distance of from 18
to more than 24 inches from the base of the plant.
Little absorption occurs in the first 2 feet of soil and this entirely from lateral
roots. The laterals are very poorly furnished with fine branches, although
at intervals of about a centimeter short branches occur not unlike those of
Astragalus. The root extremities, while very fine, are poorly branched. In
color the roots are reddish-brown, except the younger portions, which are
tan-colored. They are rather soft and more or less herbaceous, and have a
papery bark which is readily peeled off from the older ones. '
Psoralea argophylla.-This legume, which is much less conspicuous than
the preceding, forms estival societies over much of the prairie region. Plate
10, B, shows the dominant, little-branched tap-roots of several plants of this
species. A single root will be described. This had a tap (with a diameter of
8 mm.) which tapered so rapidly that at 2 feet it was less than 1 rnrn. wide,
and then pursued a vertically downward course to the water-level at a depth
of 6 feet. No large branches were given off except at a depth of 2 feet, where
a lateral occurred which was equal in diameter to the main root. This ran
quite parallel with the tap and at a distance of 4 to 6 inches from it, both
reaching approximately the same depth. -
Except for the surface foot, exceedingly fine branches, not more than an
inch in length and rarely rebranched, occurred at intervals of about 5 rnrn.
throughout the entire course of the root. At 3 inches from the tip the roots
each divided into 2 or 3 branches, all of which were branched to the first order
only. The roots are dark brown to black in color.
Baptism bracteata.-This showy legume, which forms vernal societies, has
very characteristic roots, the older portions being greenish-yellow and the
younger ones orange in color. From the base of the crown the roots spread
out obliquely and run downward, but in such a manner that the most widely
spreading were less than 18 to 24 inches horizontally away from the base of
the crown at any depth (plate 9, c). The main roots are poorly branched
THE PRAIRIES.
15
and there are scarcely any absorbing laterals in the first 2 or 3 feet of soil.
In the deeper BOil they branch irregularly but not very repeatedly, many ?f
the roots reaching the water-level at about 80 inches. It is that ill
drier lIOil they would J!t'netrate deeper. .roots near the tIps (lor
2 feet of the extremities) often form browrush, hairlike branches which may
run 12 to 18 inches without giving off laterals or, .on the other hand, may be
rofusely supplied with small branches .. Not the root-ends
up into two or three pieces, all of which are supplied WIth .laterals. In
gt"ocrw the extremities are covered with fine, short branches which extended
to the water-leveL .
THE SUBCLIMAX PRAIRIE.
During the last week in March of 1918, a number of root systems
were studied near Peru, Nebraska, at a station about 60 miles south-
caat of Lincoln. The prairies in this region are very similar in floristic
composition to those near Lincoln, as has been shown by the studies
of Thornber (1901) and others. Root systems of a number of species
were studied in a prairie area covering the exposed southeast slope of a
loess hill. A number of list quadrats which were made during the
preceding BUmmer revealed the dominance of A ndropogon furcatus
and A. 8coparius. Indeed, the striking feature of the vegetation is the
luxuriant growth of these bluestems. Andropogon furcatus extends to
the very crest of the ridge. Here the stems reach heights of 5.5 feet,
while the roots penetrate the mellow loess soil to a maximum depth of
9 foot 3 inches. This root penetration exceeds by 2 feet the maximum
depth recorded for any plant of this species in clay-loam soil.
As pointed out by Clements, this type of prairie, lying in a region
of somewhat higher rainfall, is probably sub climax. The luxuriant
growth of Andropogon upon the high ridges indicates conditions very
favorable for chaparral growth and (as indicated later) thickets of
Corylus, Rhus, and Symphoricarpos are very frequent in this grass-
land. It is not uncommon also to find seedlings of elm and oak near
the edges of these thickets. Indeed, except for fires, grazing, or other
disturbances, much of this grassland area would probably become
chaparral and forest.
Other species examined near the hilltop were Brauneria pallida
and Lygode8mia juncea. Near the foot of the slope, and where an
of clay makes the soil much harder and more compact,
specunens of Lespedeza capitata, Amorpha caneSCen8, and Ceanothus
ovatus were excavated .
is frequently the case in eastern Nebraska prairies,
WWII herb, while never abundant, occurs in such numbers as to be a
The
CODSplCUOUS component of the prairie fiora. Two plants were examined.
smaller had a tap-root 5 mIn. in diameter and reached a depth of 5.5
!:i the larger, with a diameter of 11 mIn., reached a depth of 8 feet. The
the ng tap pursues a vertically downward course, tapering very slowly. While
older parts (the first 2 to 5 feet) are more or less woody and chocolate-
brown to almost black in color, the younger parts are lighter in color and
herbaceous. Short laterals (about 9 cm. in length and mostly devoid of
branches) occur 5 to 25 mm. apart. This branching may continue to the tip,
which is about 0.3 mm. in diameter and is poorly branched. On the larger
specimen two laterals occurred (plate 11, A). The larger of these ran off at a
depth of 5 feet and at an angle of about 60
0
to a distance of 28 inches. It was
almost destitute of branches. The roots are tough and easily followed. The
coal-black streaks within the cortex and stele make their identification easy.
Although extending somewhat deeper, these specimens correspond rather
closely with those examined in the clay-loam soil at Lincoln.
Lygodesmia. juncea..-This perennial stem-xerophyte is common in prairies
throughout Nebraska and is often abundant on the crests of ridges or other
dry situations. A trench about 8 feet long and 3.5 feet wide was dug on a
slope just below a group of these plants and more than a dozen roots were
examined. The tap-root, varying from 2 to 6 mm. in diameter, may give rise
to several plants by means of short branches 2 to 8 inches below the surface
(plate 11, B).
The roots descend in an almost vertically downward course and frequently
in parallel groups only a few inches or indeed a few millimeters apart, to
distances of 15 to 20 feet or more. At these depths the tips are frequently
only 1 or 2 feet from a vertical line with the top. The roots are fleshy, very
brittle, especially after a depth of 3 or 4 feet is reached; they are from light
cream to dark brown in color and exude a white latex upon injury. The side
of the trench was dotted with drops of latex, showing the abundance of the
cut roots. This character aids greatly in recovering the broken ends of these
brittle roots. Branching occurs not at all, except for tiny laterals less than
1 mm. in diameter and an inch in length, which come off very sparingly at
almost right angles and at intervals of 6 to 12 inches. These tiny laterals are
poorly or not at all branched, the secondary branches being only 1 to 2 mm.
long. The tap narrows slowly and is frequently 2 mm. in diameter at a depth
of 16 to 18 feet. Numerous roots occurred at 18 feet in depth and one was
traced to a maximum distance of 20 feet 7 inches. Here it was still 2 mm. in
diameter. Because of the danger of caving of the mellow loess soil, deeper
excavation was abandoned. The loess was of very uniform texture through-
out and well moistened to the depth examined.
Lespedeza. ca.pitata.-This tall, conspicuous legume is common throughout
central and eastern Nebraska. It is very abundant on the lower slopes of the
loess hill, where 6 or more plants were examined. The very characteristic
light-yellow roots are much branched just below the surface, where numerous
strong laterals, 5 to 7 mm. in diameter, run off in all directions, some almost
horizontally at depths of only 3 to 6 inches. These large laterals branch very
irregularly but profusely, giving rise to numerous finely branched roots which
fill the surface soils to a distance of 2 or 3 feet on either side of the plant, when
the main branches may turn abruptly almost vertically downward (plate
11, c). In addition to these shallower roots, which compete with many of the
grasses for the water in the surface layers of soil, numerous other branches
pursue a more or less vertically downward course from the outE'et and reach
depths of 5 or 6 feet or more. These, with the vertical extremities of the
horizontal laterals, which may also reach similar depths, furnish a very exten-
sive absorbing area for the plant. The deeper roots taper rapidly, but all are
furnished with short rebranched laterals. Even the root-ends are well-
branched, but no matting was observed, as in the case of I"everal of the other
legumes. A maximum depth of 7 feet 10 inches was recorded.
THE PRAIRIES. 17
Ceanothus ovatus.-This low shrub, with its many stems 1.5 to 3 feet high,
-is quite common on prairies, especially in the eastern part of Nebraska. It
also occurs widely throughout the sandhills westward, and on the loess hill,
where the following specimen was excavated, it is rather abundant.
The plant examined was 13 years old. It had 14 stems arising from the
enlarged crown. The woody tap-root was 1.5 inches in diameter. At a depth
of 8 inches it gave off a large lateral and 2 inches below another which was
equal in diameter to the tap (2 cm.) at this depth. While the first lateral and
the tap pursued a rather vertically downward course (being, like all of the
roots, very much curved and twisted, so that at a depth of 11 feet the tap was
only 15 inches from a vertical line with the top), the second lateral at a depth
of 7 feet was 50 inches from this vertical line (plate 12, A). Also, numerous
small, repeatedly branching laterals and a few larger ones (6 mm. in diameter)
came off in the surface foot of soil and ran in rather horizontal directions for a
distance of 3 to 5 feet. In addition to these surface absorbing roots, both
short and long laterals were given off at intervals at all depths, the whole
root branching and rebranching freely. While some of the roots branched
coarsely and ended abruptly, others formed a most delicate mass of absorbing
rootlets; As a whole, the root system is well-branched, but some of the roots
at a depth of 10 or 11 feet (and where they were only 2 or 3 mm. in diameter)
ran 2 or 3 feet without giving off any branches. Numerous roots occurred
at a depth of 8 feet, several reached 12 feet, and the longest one was traced to .
a depth of 14.5 feet, where it was still 1 mm. in diameter and giving off fre-
quent threadlike laterals. The older parts of the root are woody and extremely
hard. Deeper down they lose their toughness and become very brittle, while
the smaller roots are again fairly tough. All parts of the root are character-
ized by a reddish-brown color, all but the oldest being more or less streaked
with white. They reveal a reddish color upon removal of the bark, this color
also extending into parts of the wood.
Amorpba ca.nescens.-A 7-year-old specimen, which is very representative
of others examined, was excavated near the foot of the loess hill. It gave off
11 large woody roots, 8 to 14 mm. in diameter, from the knotty crown. These
ran off at various angles from almost parallel with the hillside to almost
vertically downward (plate 12, B) where the roots are somewhat grouped,
owing to the fact that the background was only 5 feet wide. These have
v-ery few surface laterals, but those that do occur are well-branched. In the
surface 2 to 4 feet of soil relatively little absorption takes place. Andropogon
and other grasses frequently grow v-igorously between these spread-
mg Amorpha roots and doubtless suffer little competition for water. The
roots taper uniformly and at a depth of 3 feet are often still 7 mm. in diam-
eter. They frequently pursue a rather tortuous course and branch in a way
lnore or less dichotomous. Rebranching gives rise to many small laterals
only 1 to 4 mm. in diameter, which pursue a vertically downward course for
feet, giving off very few branches and tapering only slightly. Even the
tips are usually poorly branched. The lateral spread of the roots is remark-
able. One lateral reached a depth of 12 feet 10 inches and a horizontal dis-
tance of 4 feet from the base of the crown. Another was also 4 feet from a
with the crown at a depth of 12 feet 2 inches and about 5 feet from
the end of the first lateral. Similar conditions obtained on the other sides of
. eglPlant, so that a very large area of soil was penetrated by the roots of a
sm e plant.
nodules only 1 mm. in diameter occur as deep as 10 or 12 feet. On
the higher slope one plant was noted that showed much-branched root-tips
18 THE ECOLOGICAL REL4.TIONS OF BOOTS.
and nodules at a depth of 16.5 feet. The roots are smooth and chocolate-
brown in color, all being rather tough and the older parts distinctly woody.
Here again it should be noted that the plants growing in the mellow loess
reached depths 4 or 5 feet greater than those examined in the clay-loam soil
at Belmont. -
PRAIRIE ROOT SYSTEMS AND THE PRAIRIE ENVIRONMENT.
The most obvious conclusion from a consideration of these data is
the fact that prairie species are provided with well-developed, deep-
seated, and extensive root systems. Upon the basis of iot depth
the 33 species examined may be divided into three groups:
1. Shallow-rooted plants are those that seldom extend elow the
:first 2 feet of soil. These consist wholly of grasses, such as Kreleria
cristata, Stipa spartea, Elymus canadensis, Distichlis spicata, Sporob-
olus longifolius, and Aristida oligantha.
2. Plants with roots extending well below the second foot of soil but
seldom deeper than 5 feet may be grouped as intermediate in root
depth. Here belong Andropogon scoparius, A. nutans, Bouteloua gra-
cilis, Bulbilis dactyloides, Verbena stricta, H elianthus rigidus, Solidago
rigida, and Petalostemon candidus. Here also may be placed Grindelia
squarrosa, a plant which sometimes extends beyond this depth.
3. Of the plants studied, 55 per cent 'have roots which extend beyond
a depth of 5 feet; indeed, most of them to depths of from 7 to 9 feet and
a few to a maximum depth of from 13 to 20 feet or more. These may
be classed as deep-rooted species. Here belong Panicum virgatum,
Andropogon furcatus, Agropyrum repens, Solidago canadensiS, Liatris
punctata, Silphium laciniatum, Amorpha canescens, Astragalus crassi-
carpus, Psoralea tenuijlora, P. argophylla, Lygodesmia juncea, Ceanothus
ovatus, Baptisia bracteata, Lespedeza capitata, Glycyrrhiza lepidota,
Brauneria pallida, Vernonia baldwinii, and Kuhnia gZutinosa.
To understand the causes for such remarkable root development
it will be necessary for us to study the prairie environment. The
prairies of eastern Nebraska receive more moisture than most of the
great grassland area. The mean annual precipitation for Lincoln,
together with its seasonal distribution (which is of greater ecological
significance), is showil.!;in table 1. ' .
It may be seen that of the precipitation falls during the growing
season and less than of it during the three winter months.
About half of the rainfall of May, June, and July is from rains of B.n
inch or more in 24 hours. ' Such a seasonal distribution of moisture is
very favorable for the gr6:wth of grasses. Not infrequently, however,
storms occur with a rainfall exceeding 2 inches and occasionally 4 or 5
inches in a period of 24 Such storms invariably result in a high
run-off and they account for the observed deficiencieE! of mois-
ture for crops in seasons where the recorded rainfall would indicate an
'abundant supply. Drought periods of 30 or more consecutive days
between March 1 and September 30, in which precipitation to the
THE PRAIRIES. 19
amount of 0.25 inch does not occur, are not infrequent. Indeed, 16 or
17 such periods have occurred at Lincoln during the past 22 years.
The average annual snowfall is about 24 inches. "As a rule snow
covers the ground but a few days at a time after each snow storm, and
the ground is covered with snow less than half of the time even during
the months of the heaviest snowfall" (Loveland, 1912). Much of
the snow is swept by high winds intQ depressions, and thus contributes
often but little to the supply of soil moisture of the land upon which
it falls. Hence it may be seen that precipitation is only a general
indicator of conditions for plant growth. Obviously its influence upon
the distribution and seasonal activities of plants is exerted through its
power to replenish soil moisture.
TABLE. I.-Mean monthly and annual precipitation at Lincoln, Nebraska, in inchea.
Months.
Preoipi-
Months.
Precipi-
tation. tation.
Jan ....... 0.67 July . 4.01
Feb ........ 0.96 Aug ........ 3.72
Mar ........ 1.26 Sept ...... 2.91
Apr ........ 2.51 Oot ......... 1.94
May ....... 4.39 Nov ........ 0.94
June ....... 4.43 Dec ........ 0.85
Annual . 28.59
The fertile, dark-colored prairie soil of the region is of the type com-
monly called loess, much of which, however, is confounded with glacia1,
drift. "The loess' covers the hills and valleys alike to a depth of from
20 to 100 feet, being much thicker than this in places and much thinner
in others. Throughout the first 100 miles westward from the Missouri
it is underlain by Kansan till" (Alway,1916). "The uniformity in the
physical properties, recognized as characterizing the material of the
loess, should tend to produce, under uniform climatic conditions, soils
uniform in chemical properties." The water-holding capacity of the
~ c e foot of soil is about 60 to 70 per cent of its dry weight, while the
moisture equivalent and wilting coefficient are 25 per cent and 13.5
per cent respectively} .
Studies of the water-content of upland' prairie sojIs have been
carried on for more than two seasons. Table 2 . gives the results of
water-content determinations during the growing season of 1916.
The minus sign indicates water non-available for plant growth. .
A glance at these results shows that at four different periods no
water was available for growth at a depth of 4 inches, while during
late August the same condition obtained for the 4 to 12 inch layer.
Unfortunately, deeper soil samples were not obtained, but during the
follOwing seasons samples were taken at irregular intervals to a depth
of 5 feet. These data are shown in table 3.
~ The writer is indebted to Dr. L. J. Briggs, of the Bureau of Plant Industry of the U. S
...... partment of Agriculture, for determinations of moisture equivalents- and wilting coefficients.
20
TABLE 2.
In tables showing water-content it will be understood that the figure columns represent
J'erCentages.
Depth of samples.
Depth of samples.
Date.
o to 4 inches. 4 to 12 inches.
Date.
o to 4 inchea. 4 to 12 inches.
Wilting Wilting
Wilting Wilting
coefficient 12. coefficient 12.7.
coefficient 12. coefficient 12.7.
May 21. .. 19.5 17.3 July 31. ...
- 0.8 0.1
June 5 ... - 2.5 4.8 Aug. 7 .... 9.5 8.8
June 10 ... - 3.5 3.8 Aug. 15 .. 2 O ~ O 13.8
June 17 .... 3.0 9.1 Aug. 22 ....
- 2.2 -2.7
June 26 .... 1 6 ~ 5 9.5 Aug. 28 ....
- 1.7 3.8
July 1 .... 1.3 5.5 Sept. 4 .... 13.1 8.8
July 8 .... - 2.4 2.0 Sept. 12 .... 13.5 5.3
July 17 . 9.0 2.8 Sept. 19 .... 2.0 1.8
July 24 ... - 1.2 0.3
.
. TABLE 3.-At1ailable toater-content oj the prairie BOil during 1917.
The minus sign indicates water non-available for plant growth.
Depth of Depth of Depth of Depth of Depth of Depth of
sample, sample, sample, sample, sample, sample,
Date.
Oto6 6 to 12 1to2 2to3 3to4 4too
inches. inches. feet. feet. feet. feet.
Wilting Wilting Wilting Wilting Wilting Wilting
coefficient, coefficient, coefficient, coefficient, coefficient, coefficient,
13.4. 13.4. 15.4. 14.5. 16.1. 16.1.
Apr. 3 ............ 4.3 4.3 -2.6 -1.6 -2.9 -0.4
May 6 .... -2.9 6.7 2.2 -1.4 .......... ...........
June 19 ........ 4.9 9.8 8.1 8.4 6.1 4.3
July 9 ............ -1.9 2.0 .......... .......... .......... ..........
July 16 ............ -1.7 0.7 2.1 1.7 .......... ..........
July 23 ............ -3.1 -1.7 .......... .......... .......... ..........
July 30 ........... -5.3 -2.3 -4.0 .......... .......... ..........
Aug. 6 ............ 1.6 -1.3 .......... .......... .......... ..........
Aug. 13 ............ 7.0 0.5 .......... .......... .......... ..........
Aug. 28 ... 2.1 1.7 -2.1 -1.3 -4.5 -1.9
July 24 (1916) ...... .......... ............ -0.9 2.3 . .......... ..........
Jan. 5 (1918) ...... 3.3 1.9 -0.9 -0.7 0.6 2.1
Mar. 26 (1918) ...... 7.2 7.4 1.4 -3.9 -4.2 -2.8
On May 6 and again throughout the whole of July, no water was
available in the first 6 inches of soil. During late July and the first
week of August soil moisture was depleted to a point below the wilting
coefficient for a depth of 12 inches. On April 3 and again on August 28
no water was available at a depth of 1 to 5 feet. These results .are
rather surprising, but an exa.miD8,tion of the rainfall record shows that
during 1916 the precipitation was 5.5 inches below normal, while during
1917 it fell to 6.5 inches below. The dryness of the soil during 1916-17
was shown by an abnormal amount of winter-killing of trees and
shrubs. During July the rainfall was only 0.56 inch instead of the
-normal 4 inches. The determinations on July 24 (1916) at 2 and 3
feet respectively show that the soil was very dry. It is interesting to
note in this connection that similar results were obtained in the
THE PRAIRIES. 21
prairies of southeastern Washington, where during 1914 the water-
content to a depth of 4 feet was reduced to the non.,.available point
(Weaver, 1915: 233-235). .
The available water-content in the loess soils supporting the sub-
climax prairie vegetation was not only higher but much more con-
stant. These data are shown in tables 4 and 5. Here also are included
determinations from an adjacent thicket of CoryZus americana. A
comparison of these results is made on page 30.
TABLE4.-Available water-content in prairie and ahrt.tb communities near Peru, Nebraska,
during April to September 1917 and May 1918.
1
The minus sign indicates water non-available for plant growth.
Depth, 0 to 6 inches. Depth, 6 to 12 inches. Depth, f to 2 feet. Depth, 2 to 3 feet.
Wilting coefficients: Wilting coefficients: Wilting coefficients: Wilting coefficients:
Prairie 11.7; scrub 13.8. Prairie 10.9; scrub 12.3. Prairie 10.4; scrub 12.5. Prairie 9.9; scrub 11.3.
Date. Prairie. Scrub. Date. Prairie. Scrub. Date. Pram e. Scrub. Date. Prairie. Scrub.
1917 1917 1917 1917
Apr. 5 13.4 21.4 Apr. 5 12.2 18.0 Apr. 5 8.5 12.1 Apr. 5 5.0 8.0
May 15 13.0 14.7 May 15 12.0 14.1 May 15 10.5 11.7 May 15 8.1 ......
June 30 13.9 20.8 June 30 11.8 17.1 June 30 12.3 13.1 June 30 12.6 12.9
July 9 6.5 3.9 July 9 7.3 7.4 July 9 .. ...... July 9 .. . .....
July 14 9.5 11.5 July 14 7.4 5.2 July 14 ........ ...... July 14 . ....... ......
July 24 0.1 4.5 July 24 2.5 3.4 July 24 3.1 3.0 July 24 5.1 5.5
July 28 -1.4 -1.7 July 28 0.4 -0.4 July 28 2.1 1.2 July 28 4.5 4.4
Aug. 6 4.0 6.6 Ailg. 6 -0.5 2.6 Aug. 6 -1.0 0.7 Aug. 6 1.9 2.4
Aug. 11 8.4 12.6 Aug. 11 1.5 5.7 Aug. 11 ........ ...... Aug. 11 . ....... ......
Aug. 18 2.3 7.6 Aug. 18 1.9 1.5 Aug. 18 ........ ...... Aug. 18 . ....... ......
Aug. 25 1.3 6.4 Aug. 25 0.2 1.8 Aug. 25 0.6 -0.1 Aug. 25 0.9 '1.2
Sept. 8 6.2 10.2 Sept. 8 -0.8 2.4 Sept. 8 0.8 -0.4 Sept. 8 1.6 -1.0
Sept. 22 -1.2 -1.6 Sept. 22 -1.7 0.5 Sept. 22 -1.0 -0.7 Sept. 22 0.1 0.5
1918 1918 1918 1918
May.16 13.8 16.3 May 16 15.0 18.3 May 16 12.5 16.5 May 16 12.7 16.5
1 The writer is indebted to Prof. F. C. Jean for the major portion of the data in these tables.
The greater water-content of these mellow loess soils compared with
those of the prairie near Lincoln gives us a clue to the much more
luxuriant growth of certain prairie and shrub species. There is also a
great difference in depth of root-penetration, plants like Andropogon
Jurcatus and Brauneria paZZida being much deeper-rooted in loess soil.
However, vegetation is not only an expression of present conditions,
but also to a greater extent a record of conditions that have obtained
during a period of years, and the record is not likely to be alterp.d
greatly in a year or two in which conditions may depart from the
normal. The preceding pages show that many prairie plants absorb
moisture well beyond a depth of 5 feet, while soil-moisture extends
many feet beyond the greatest root depth. In excavating root systems,
during September to December 1917, the soil below 5 feet was found
almost invariably to be quite moist. Some of the root systems were
excavated on bench-lands in the Salt Creek basin, where the alluvial
soils are somewhat different from those described. Such differences
Were noted in discussing these species.
Just as the possible growth of the aerial parts of plants is affected
by the extent of the development of the root system, conversely the
environmental conditions to which the aerial parts are subjected,
especially as concerns their water relations, must reflect themselves
in the root development. Therefore, it will be instructive to consider
briefly the above-ground environment.
TABLE 5.-Total wateT-content oj soil during 1917. TABLE 6.
3 to 4 feet. 4 to 5 feet.
Date.
Month. Temp. Month. Temp.
Prairie. Scrub. Prairie. Scrub.
OF. OF.
1917 p. ct.
Apr. 5. 15.9 16.6 13.0 16.1
June 30. 20.5 24.7 21.4 24.9
Aug. 6 . 15.5 15.4 16.9 16.7
Sept. 22. 12.2 12.1 11.9 13.0
1915
May 16. 13.1 15.0 15.4 15.1
Jan . 22.6 Aug . 75.0
Feb ... 24.3 Sept .. 66.6
Mar .... 37.4 Oct M.6
Apr .... 51.5 Nov . 40.4
May ... 61.7 Dec . 2S.5
June .. 71.2
July ... 76.4 Aver .. 5O.S
The mean monthly temperature at Lincoln, Nebraska, for a period of
more than 30 years is shown in table 6.
The season without killing frosts usually extends from the first day
of May into the first week of October, but frosts have occurred as late
as the last week of May and as early as the second week of September.
Figure 4 shows the average daily temperatures obtained during the
growing season of 1916, as recorded by a hygrothermograph placed
in an appropriate shelter in the prairie at a height of 17 cm. The
weekly means were obtained by drawing a horizontal line through the
weekly record sheet in such a manner that the total area above this
line included by the graph was equal to the total area below the line.
M JUNE JULY A U ~ .
~
40
35
30
25
20
15
10
M .JUNE
1\
I \
IY
"i \
1/\
I".
V
.JULV AUG. SEP.
~ \
1\\
1""/ \
,..
~ \/'1.. /
J
I'"
FIG. 4.-Graphsshowingtheaveragedaily FIG. 5.-Graphs showing the aver-
temperature (heavy line) and humid- age daily evaporation rates in
ity (light line) in the prairie during the prairie during 1916 (heavy
1 9 1 ~ . line) and 1917 (light line).,
The areas were determined by the aid of a planimeter. In this inter-
pretation both temperature (or humidity) and the time factor are
taken into consideration. This figure also gives a record of the mean
THE PRAIRIES. 23
weekly humidity of the air during this same period. Not infrequently
during the long summer afternoons the humidity falls to 20 per cent
or less, while the temperature may reach 95 to 100 F.
During the growing season the prevailing winds are from the south
or southeast. A standard anemometer placed in the prairie on a gentle
southwest slope and at a height of 0.5 meter recorded 4,905 miles of
wind from July 13 to September 19, 1916, an average of 72 miles per
day. A similar instrument placed at the same height above the soil
surface, but on a hill top in the same pramearea, showed an average
daily wind movement of 122 miles per day.
The factors of high temperature, low humidity, and wind movement
combine to increase the evaporating power of the air. Livingston's
porous-cup atmometers, fitted with non-absorbing mountings, were
operated in the usual manner in duplicate and at a height of about 17
cm. above the soil surface during 1916 and 1917. The combined
evaporation records from two prairie stations, reduced to the readings
of the standard cup, are shown in figure 5. During the last week in
July 1916, when the mean daily temperature was 81.4 F. and the
mean relative humidity 57.4 per cent, the daily evaporation was 38
c.c. During this period no soil moisture was available in the first foot
of soil. The average daily evaporation throughout this whole summer
was 21.6 c.c. Atmometers exposed at a height of 0.5 meter during the
last 10 days in July gave losses over 50 per cent greater than similar .
instruments placed at the usual height of 17 to 23 cm. In the prairies
near Peru, Nebraska, the average daily evaporation from May 15 to
" September 22,1917, was 20.9 per cent.
Summarizing briefly, we find that prairie plants of this region grow
under semi-arid climatic conditions in which the supply of water is the
chief limiting factor of plant growth. During certain portions of the
growing season extremely xerophytic conditions are brought to bear
upon the vegetation. It has been shown that the water-content of the
soil is reduced to the non-available point to a depth of 4 or 5 feet at
least during certain years and often at times when "the evaporating
power of'the air is very high (38 c ~ c . daily).
In response to these environmental conditions many species have
developed surprisingly extensive root systems. In fact, all of the
dicotyledons examined, as well as many of the grasses, extend their
root systems to depths of from 3 to 7 feet or more. For just as the
evaporating power of the air and the nature of the transpiring organs
determine the water-loss of plants, likewise the soil water and the nature
of the root systems determine the supply. These findings of great root
depths, which are correlated with deep soil moisture, bear out Can-
non's suggestion of the probability that the longest or most deeply
penetrating roots are found, not in deserts, but where there is consider-
able rainfall, and where the penetration of rain is considerable and the
water-table relatively deep.
The roots of prairie plants are grouped into more or less definite
absorbing layers, many of the deeper-rooted species having few or
no absorbing roots in the first few feet of soil. This layering of the
roots reduces competition and permits the growth of a larger number
of species. Any of the shallow-rooted grasses extract water from the
soil layers quite above those from which Andropogon furcatm or
Solidago rigida draw their supply, while still deeper soil strata furnish
moisture for such deep-seated species as Panicum virgatum, Silphium
laciniatum, and Kuhnia glutinosa. It should be pointed out here that
prairie plants very seldom show the widely spreading and superficial
laterals so characteristic of many plains species. The cause seems to
lie in the greater water-content of the deeper soil in the prairie.
Root variations of prairie plants may be due to a number of factors,
among which the water-content of the soil and its penetrability prob-
ably stand first in importance, at least in non-alkaline soils. Although
the problem of the causes of root variation is one which must be solved
under control conditions where one factor can be varied at a time,
considerable evidence has been found in the field that both soil mois-
ture and soil texture profoundly influence root distribution.
The effect of compact soils upon root penetration is shown not only
by the tortuous courses pursued by roots and the distortions of the
roots themselves, but also by their modified branching. Several
species, including grasses, were found to penetrate from 2 to 3 feet
deeper in deep loam soil than in the same type of sQil when it was
underlaid with a hard clay subsoil. In soils with a subsoil of alternating
layers of sand and clay a striking distribution of lateral roots was
observed. These often occurred abundantly near the bottom of the
sandy layers and in the clay strata where the latter acting as a rather
impervious layer had retained much soil moisture.
It is interesting to note that species of the same genus may have
an entirely different type of root system.. Liatris punctata has a
deep tap-root, while the multitudinous roots arising from the corm of
L. scarWsa are fibrous. Likewise, the roots of Andropogon .scoparim
are as different from those of A. furcatm as are the roots of KtBleria
cristata from those of Stipa spartea. Indeed, the roots studied are
remarkable for their individuality. The roots of each species, because
of peculiarities of form, branching habit, position in the soil, texture,
color, odor, or taste, can be easily distinguished, and these distinguish-
, ing have o.ften proved useful in some
.' cases It has be'enposslblV to reconstruct ,uCCesSlOna.1 from the
, record left in the soil by 'partially preservrd roots and rJUzomes.
THE CHAPARRAL COMMUNITY. 25
II. THE CHAPARRAL COMMUNITY.
Between the great Ohio-Missouri deciduoustforest complex on the
east and the prairies to the westward, there extends a chaparra
community. Indeed, this is more or less continuous from Canadl
to Texas. In this shrub community, tongues of which extend far into
the sub climax prairie, species of Symphoricarpos, Rhus, Corylus, and
Rosa play the rOle of dominants. Indeed, all but the latter form
thickets of greater or less extent in the moister places throughout much
of the prairie community or occur as more or less isolated clumps or
individuals held in check only by the severe root competition of the
prairie species (plate 16, B). They rapidly spring into dominance,
suppress the grasses, and form centers for further outward invasion,
when soil-moisture conditions are even slightly increased above the
normal. This may be due in the first instance to the loosening of the
soil by burrowing animals, the building of a fence, or other disturbance
of the sod.
In order to understand more clearly the nature of the competition
between scrub and grassland, as well as to determine more exactly
environmental conditions indicated by these phyads, a number of root
systems were examined along the loess hills of the Missouri River near
Peru, Nebraska.
SYmphoricarpos vulgaris.-This species, together with its western ecological
equivalent, S. occidentalis, is a very conspicuous and important shrub of the
chaparral community. Growing in clumps to a height of only 2 to 4 feet,. its
shade is frequently so dense as to exclude even the very tolerant Poa pra-
temia. In a well-established shrub area the latter almost invariably occurs
between the individual clumps.
A long trench was dug at the edge of such a thicket and the roots of numer-
ous plants were examined. The larger roots arise mostly from near the base
of the erect shoots, but numerous smaller ones occur, especially all along the
underground stems (plate 13, A). The maximum depth to which the roots
penetrate in the loess soil is only 65 inches. Although the trench was sunk to
a depth of about 8 feet and a part of the soil underlying the thicket at this
depth was removed, no roots of Symphoricarp08 were found below the 65-inch
level. Indeed, except for a few plants of Rhus, competition for light above
ground was so intense that no other plants were present and the deeper soil
was free from roots of any kind. But lack of linear extent is amply recom-
pensed by a wonderfully well-developed absorbing surface. Perhaps the roots
of no other plant examined, with the exception of certain surface-feeding
grasses, occupy the soil more thoroughly than does the delicate network of the
root branches of this shrub.
The larger roots vary from 3 to 7 mm. in diameter. While they may come
off vertically, usually they pursue an oblique direction for some distance
(1 to 3 feet) before turning downward (plate 13, A). These reddish-brown,
tough, woody roots taper uniformly, frequently giving off large branches and a
network of finer ones, beginning just beneath the surface of the soil. Indeed,
the roots are profusely and minutely branched and rebranched throughout,
the ultimate branches being almost microscopic in size. The laterals vary
in length from those only an inch long to others extending a distance of 2 or
more feet, sometimes almost at right angles. The last foot or 18 inches of the
root is usually threadlike but extremely well branched. Above the level of
65 inches the soil is well filled by great numbers of these fine root-endings.
Rhus glabra.-A trench 10 feet long was dug just within a thicket of sumac
which, according to ring counts, had been in possession of the area for at least
12 years. Practically all herbaceous species had disappeared except Poa
pratensis. A brief examination showed the soil to be filled with a network of
rhizomes from which numerous roots ascended more or less vertically and
ended at the surface of the soil in a brush-like mass of branches. Further
study revealed the really wonderful complexity of these underground parts
as well as the great absorbing area of this xero-mesophytic shrub (plate
14, A, B). Roots were examined in three different situations. Although they
differ in detail.3, the following is a typical example:
A tap-root 3 cm. in diameter and 12 or 13 years old pursued a downward
course at an angle of about 40 from the vertical to a depth of 22 inches. Here
it turned off at an angle of 30 from the horizontal and ran a distance of 2
feet, at which length (48 inches) it was only 1.5 cm. in diameter and 37 inches
deep. Here it forked into two equal branches, one (a) running almost hori-
zontally to a distance of 68 inches and at a depth of 3 feet, while the other (b)
turned off at an angle of 40 from the first and, after running rather obliquely
downward for a distance of 15 inches, gave off,two horizontal laterals 3 and 6
rom. in diameter respectively, while the main root turned vertically upward.
It ran in this direction for a distance of 8 inches, when it divided into 3 equal
branches. Two of these were each 6 rom. in diameter and tapered gradually
till they reached the surface, 26 inches above, and at a horizontal distance of
26 inches beyond the vertical root from which they branched. They ended
about 2 feet apart in an extremely well-branched and rebranched network
of fine rootlets. These root termini were 70 inches in a horizontal line from
the beginning of the tap.
The third branch from the upright portion of the root turned off horizon-
tally and ran a distance of 8 inches, when it again turned at right angles and
ran horizontally in another direction (away from the main tap) 16 inches,
finally turning downward at right angles and descending to a depth of 80
inches. The tip, which ended in a much-branched network of almost hair-
like terminals, was located at a horizontal distance of 68 inches from the .
beginning of the tap. The two 3 and 6 rom. laterals of root (b) ran off hori-
zontally, but in a direction opposite from the others, finally ascending to the
surface several feet from the other roots. Branch (a), which was partly cut
off in digging the trench, also sent off numerous roots to the surface as well as
some into the deeper soil. The surface soil is filled with rootlets with such a
network of absorbing terminals that competition with the grasses for soil
moisture must be very severe. Nor is the main absorbing area confined to the
surface. Well-branched laterals, both large and small, arise at all depths, and
even the deeper soil is rather thoroughly occupied. The well-branched root
system, together with the large area it occupies, must afford abundant mois-
ture, even in fairly dry soil.
While all of the main roots examined in two separate thickets showed the
oblique to horizontal course with a depth never exceeding 80 inches, one root
of a Rhus plant growing in the Symphoricarpos thicket was 0.5 inch in diam-
eter and pursued a vertically downward course to a depth of over 90 inches.--
The roots are characterized by a thick cortex which exudes a white latex when
injured, and upon removal reveals the small, woody, glistening white stele.
The younger roots vary in color from brown to nearly black.
Corylus americana.-This shrub does not get so far into the prairies from
the bordering eastern forests where it forms a layer as do the preceding, and
in this respect shows its less xerophytic habit. But in many situations, from
Minnesota to Kansas, thickets of Corylus come into direct competition with
prairie vegetation.
Corylus spreads by means of large woody underground parts, as shown in
plate 15, c. These run at a distance of only 4 to 6 inches under the surface
and give rise at intervals to numerous erect shoots from 2 to 7 feet high and
also to multitudinous roots, some of which are more than 1.5 cm. in diameter.
Some of the smaller roots run vertically toward the surface and branch pro-
fusely into very fine ultimate branchlets, for Corylus, like the preceding
shrubs, is a strong competitor of the grasses for moisture in the surface soil.
Indeed, the surface foot or two of soil is well filled with absorbing rootlets, the
longer roots being only fairly well supplied with laterals to a depth of 10 or 11
feet. These larger roots, as shown in plate 15, A, either pursue a rather verti-
cally downward course, or extend obliquely for a distance of 1 or 2 feet or
more and then turn downward. Roots were traced to depths of 10, 10.5, and
11.5 feet respectively. They are dark brown in color, very woody, with a very
thin cortex, and frequently branch rather dichotomously, although the
branches are seldom equal in size. The branches are often very coarse and
sometimes run in parallel groups several inches without giving rise to absorb-
ing rootlets. The main root-ends are not very well supplied with fine branches.
Thus it may be seen that while Corylus penetrates to greater depths than
Rhus or Symphoricarpos, its absorbing system is hardly developed to such a
high degree.
Vitis vulpinaA-At a depth of over :w. feet in the Rhus thicket a root of
Vitia was encountered. It was 18 rom. in diameter and ran horizontally
across the trench. It gave off numerous branches, some of which reached
depths of 12 feet. Its branching is not unlike that of the larger roots of the
hazel. In the Corylus thicket, roots of grape were traced to depths of 13 feet
3 inches and 13 feet 6 inches respectively. The diameter of roots near the
surface was abOutiCiii.; at 10 feet it was 2 and 3 mm. respectively. Another
underground part ran horizontally at a depth of 2 feet for a distance of more
than 6 to 8 feet. Such an enormoup absorbing area as is possessed by Vitia
gives us the clue to its ability to have a leaf area not unlike that of many trees
andyet be able to live often in apparently rather dry situations.
Rosa arkansana.-This shrub is widely distributed throughout the scrub
and prairie areas. Although usually held in check in the dense grassland sod,
except where local disturbances have favored its development, in the sub-
climax grassland it frequently forms dense brush over the less-exposed slopes.
In the sandhills westward it again assumes an important r61e, areas several
acres in extent frequently being controlled by this species.
As shown in plate 16, A, this plant propagates by rhizomes. This parent
plant had given rise to 5 distinct clumps of stems, the youngest of which was
5 years old and more than 5 feet from the oldest. Fragments of other hori-
zontal parts, which had not yet developed shoots, may be seen. It may also
be noticed that the plant next to the parent is the only one that has developed
a tap-root of its own. Indeed, the other roots arising from the horizontal
portion ran obliquely upward, rather horizontally or, at most, only very
obliquely downward, and none reached any considerable depth when com-
pared with the taps from the older plants. The tap-root from the second
~ l a n t reached a depth of 15 feet 2 inches; the main tap pursued a nearly ver-
tlCally downward course to a depth of 21 feet 2 inches.
It shoulq be noted here that these roses grew about midway up a southeast
slope. Here the loess soil was intermediate between that described for
Lygodesmia near the crest and for Amorpha near the foot of the hill. Beyond
a depth of 10 feet the soil was quite compact.
Plate 15, B, illustrates well the paucity of large branches. The lateral
spread of any branch measured horizontally from the base of the crown did
not exceed 4 feet. Although many fine branchlets occurred along the course
of these main roots and extended off laterally for distances of 6 to 18 inches,
still other portions were quite free from branches. The breaking up of the
larger roots near their extremities into numerous long, slender, often more or
less parallel rootlets is well shown on the root in the figure, which ends at a
depth of about 6 feet.
The older woody roots can easily be identified by scraping off the outer
black part of the thin cortex, which then reveals a bright red color, while
similar treatment of the younger roots shows their orange color.
SHRUB ROOT SYSTEMS AND THE SHRUB ENVIRONMENT.
From the preceding descriptions it may be seen that the members
of the chaparral community of the tension zone are all supplied with
splendid absorbing systems, which are somewhat variable as to depth,
but all of which are deep-seated. In addition, all have excellent
-methods of vegetative propagation.
The r6le played by these shrubs in modifying prairie conditions to
such an extent that the habitat becomes rather favorable to the growth
of the more xerophytic trees, such as bur oak, should not be overlooked.
All are well adapted, either by means of above-ground or under-
ground stems or root offshoots, to slowly but successfully invade the
prairie sod. For example, the whole loess hill, where these shrubs were
excavated, would probably be covered by a chaparral complex except
for repeated fires which are more detrimental to shrubs than to grasses
and most other prairie species. The proof of this statement lies not
only in the presence even over much of the more exposed hillside of a
potential scrub mictium, but also in the fact that Rhus, Symphori-
carp08, and other shrubs dominate over other portions of the same
ridge where soil conditions are identical but where fires have not been
permitted to run. '
Synphoricarpo8 extends its area by migrating by above-ground stems
as well as by those below the surface. This mechanism of invasion ;
is well illustrated in plate 13, B. While the above-ground stems furnish
the more rapid method of migration, they frequently fail to become
rooted in the prairie sod. They then become greatly attenuated and
soon die. The under-ground stems are more certain of establishment.
Once established, this shrub rapidly reacts upon the habitat by
increasing the water-content both of air and soil, and also by modifying
the texture and composition of the latter. The presence of the shrubs
decreases wind movement and their shade reduces not only the light
but the temperature as well. This reacts favorably upon the humidity
and also decreases the evaporation from the soil surface. The reduc-
tion of the evaporating power of the air within a Symphoricarpo8 area
at a height of 17 cm. as compared with that 2 meters beyond in the
prairie is well illustrated by the following data, giving the average
daily evaporation in a clump of Symphoricarpo8 and in the adjacent
grassland, both near the crest of a loess hill:
Date. Prairie. Scrub.
c. c. c. c.
July 1 to 7, 1918 ..... 33.1 25.6
_ July 29 to Aug. 5, 1918 .. 46.0 33.8
,4ug. 26 to Sept. 4, 1918. 18.0 14.6
.--
Not only is the evaporating power of the air lowered by the presence
of the scrub, but among its stems drifts of snow find lodgement and
upon melting add considerably to the moisture content of the soil.
The run-off is greatly reduced as a result of the rich mulch of fallen
leaves and large quantities of wind-transported plant debris lodged
among its stems. Only a few years are required, owing to the more
favorable conditions for the formation of humus under the shrubs, to
fill the former prairie soil with a rich humus mulch not unlike that of
the woodland. Thus by cumulative favorable reactions the scrub
is frequently able to extend its area of dominance and also to prepare
a fine nursery for the seedlings of trees, thus initiating a new stage in
the normal succession. Repeated determinations of water-content to
a depth of 3 feet show that the thicket soil is moister than that of the
prairie.
The reactions of Rhus upon the habitat are similar to those de-
scribed for Symphoricarpo8. Being a taller shrub, its effect upon the
bordering grassland is more pronounced. It invades the prairie by
means of rhizomes. These have been traced at a depth of only 4 to
8 inches to a maximum distance of over 20 feet from the edge of a
thicket, their course being marked by the presence of erect shoots.
The evaporating power of the air is greatly reduced under Rhu8 as
compared with that in the prairie, as is shown by the following experi-
ment: From May until September, 1916, non-absorbing atmometers
placed just 6 meters within the sumac thicket gave evaporation losses
averaging about 10 c.c. per day as compared with average daily losses
of 22 C.c. from similar instruments 24 meters outside the thicket in the
prairie. Likewise, the water-content of the soil at the scrub station
as compared with that of the grassland was found to average about
6 per cent higher throughout the season to a depth of 10 inches. The
deeper layers of soil to 5 feet were invariably much moister in the
sumac community.
The profound effect of a growth of hazel upon modifying prairie con-
ditions is well illustrated by a series of evaporation and soil-moisture
readings obtained on a flat hilltop in the edge of a hazel thicket and a
few meters beyond in the prairie. The evaporating power of the air
measured during the growing season of 1916 and 1917 was about 50
per cent less in the Corylus thicket. On the other hand, the water-
content of the soil, as in the case of the other shrub communities, was
found to be somewhat higher. .
During 1917 another series of evaporation readings and sob-moisture
determinations was made at Peru in the loess soil. The atmometers
gave an average daily loss of 20.9 c .. c. in a prairie on a southeast slope,
while in a Corylu8 thicket on a northwest slope the loss was only 14.4
c.c. daily. This ratio was maintained almost throughout the summer
and at no time were the losses in the shrub' as great as those in the
prairie. That the water-content of the soil in the shrub was usually
much higher than that in the prairie_may readily be seen by an exami-
nation of tables 4 and 5 on pages 21 and 22.
THE PRAIRIES OF THE PACIFIC NORTHWEST 31
III. THE PRAIRIES OF THE PACIFIC NORTHWEST.
The prairies of southeastern Washington and adjacent Idaho,
where the following studies were carried on, represent an extreme
westward extension of the great grassland formation lyigg-east--ef- the
Rocky Mountains.- Agropyrum spicatum, Festuca ovina ingrata,
Kreleria cristata, and Poa sandbergii are dominants. Stipa is entirely
absent, as are also the late-blooniing grasses, such as Andropogon,
Bouteloua, and others of the eastern prairie. The absence of these
late-maturing grasses may be accounted for by the peculiar distribu-
tion of the precipitation. Only about one-fourth of the annual 21
inches of moisture falls during the growing season. Except for the
extremely retentive silt-loam soil, the region would be almost a desert.
This is an extremely important fact to keep in mind while consider-
ing root systems. The important r6le played in the eastern prairies
by Andropogon scoparius is here taken by Agropyrum spicatum, its
ecological representative. Like Andropogon, it presents the bunch
habit in drier soils, but becomes a sod-former with well-developed
-rhizomes under more favorable moisture conditions. Eastward, these
prairies adjoin the main, area of grassland lying east of the Rocky
Mountains. At Missoula, Montana, for example, the Agropyrum-
Festuca community of Washington and Idaho (Weaver, 1917) meets
and intermingles with such eastern prairie species as Stipa comata,
Aster multiflorus, etc., while Agropyrum spicatum, Kreleria cristata,
Solidago missouriensis, Achillea mille!oZium, and others occur through-
out both regions.
The writer has presented elsewhere (1917) a detailed description of
the extent, successional relations, and floristic composition of these
western prairies. A glance at plate 17, Band c, may give some impres-
sion as to their general character. Hence we will proceed at once to a
discussion of root distribution.
THE ROOT SYSTEMS OF THE GRASSES.
Over 60 individuals of the four dominant grasses were excavated
and examined. Three, Kreleria cristata, Poa sandbergii, and Festuca
ovina ingrata, are shallow-rooted, the bulk of the absorbing system
lying above the IS-inch level, while Agropyrum spicatum penetrates to
a maximum depth of 4 feet 10 inches.
Agropyrum spicatum..-This is the dominant bunchgrass in eastern Wash-
ington. It has its best development westward of the high upland prairies
of extreme eastern Washington and along the rim-rock through the eastern
part. The bunches are often 10 inches in diameter and reach a height of over
3 feet. The plant blossoms in June and dries out in early July, only to take
on renewed growth after the autumn rains and to remain green all winter.
This grass has coarser roots than any of the other three important native
grasses. _ These coarse, fibrous roots have many short laterals. Some of the
feet
8
61- Il/Jn}l\\ ,,\llml\\.. 'J I '.J{\' ,1 1/11 I (l \'1\ Id' I v' J/ Arm\()) Jo..-<:-::n "-L.:..:..r1\:, q I )..' '( 11\ ' l l " I ~ O m ' -I
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6
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FIG. 6.-Schematic bisect showing the root and stem relations of important prairie plants. This and figures 7 and 8 were drawn from photo-
graphs and data obtained by the excavation and examination of 325 root systems of these 18 species: h, Hieracium Bcouleri;
k, K(JJleria crUtata; b, Balsamorhiza Bagittata; J, FeBtuca ovina ingrataj g, Geranium viBcoBiBBimumj p, Poa aandbergii; ho, Hoorebekia
racemoBaj po, PotentiUa blaBchkeana.
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THE PRAIRIES OF THE PACIFIC NORTHWEST. 33
roots reach a depth of 4 feet 10 inches, although on an average 4 feet 2 inches
was the greatest depth attained.
Festuca ovina. ingrata.-The blue bunchgrass ranks m importance with
Agropyrum on the well-developed high prairies west of the foothills of the
Bitterroot Mountains between Spokane, Washington, and Lewiston, Idaho.
Because of its abundance the very appropriate name Palouse (Fr. pelouse, a
land clothed with a short, thick growth of herbage) was early applied to this
region. The whole plant dries out considerably by the middle of July, but the
autumn rains revive it and it is green.throughout the rest of the year.
Festuca ovina has a great mass of jet-black roots which occupy the soil
thoroughly from the surface to a depth of about 18 inches, below which depth
relatively few roots extend. None of the roots are over 1 mm. in diameter.
They branch profusely to the third order mostly, and the laterals are usually
less than an inch in length. This branching continues to the very tip, and
there the laterals are usually longer. The longest root found was 3 feet 3
inches, and the average length Wail 2 feet for the deepest roots, but the great
bulk of these roots were less than 18 inches long.
Poa. sandbergii.-This species grows in small tufts, usually only from 0.5
to 1.5 inches wide, puts out new roots when the fall rains begin, grows through-
out the winter and spring, and evades drought by flowering late in Mayor in
early June and remaining dormant the rest of the growing season.
Poo has smaller roots than Festucaj they are more profusely branched,
and the fine, short laterals are more numerous, smaller, and much more
branched. The creamy-white roots spread laterally 3 to 5 inches and occupy
thoroughly the first few inches of soil, relatively few extending below a depth
of 8 inches, and none was found beyond 13 inches. The root branches are
longer and more numerous at the tip than are those of Festuca. The average
maximum depth was found to be 10 inches.
Kreleria. cristata.-This is also a dominant bunchgrass on the p r a i r i ~ of
eastern Washington. It flowers in late June or early July, and like Poa
remains dormant until revived by the autumn rains. The roots resemble
those of Agropyrum, but taper faster and have finer laterals which branch
mostly to the third order. These laterals, like those of the shallow-rooted
Poo, are more numerous than in Agropyrum. The deepest root found was at
28 inches, and 15 inches was determined as the average maximum depth.
The shallow root habit of the t ~ last species as compared with the deeper-
root habit of Agropyrum and certain' dicotyledonous plants is shown graph-
ically in figures 6 and 7;
THE ROOT SYSTEMS OF OTHER PRAIRIE SPECIES.
Since the root systems of 21 of the most important non-grassy
prairie species have been described in detail in a former publication
(1915), it will be necessary here only to present a snmma.ry statement,
so that it may be possible to compare their root systems directly with
the plants of the eastern prairies and later with those of the plains.
Such a comparison will help us to more clearly evaluate the response
to the conditions under which the several plant communities grow.
Lupinus ornatus.-This plant and L. leucophyllus are the two species of
lupines most widely distributed on the high prairies of eastern Washington.
Both form extensive summer societies, the former on the drier slopes and
ridges, the latter on moist hillsides and in the valleys. Several of the 24 root
feet
4
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FIG. 7.-Schematic bisect: 8, SieverBia ciliata; 10, Wyethia amplezicami8; l', Lupin'U8 Ze'UCOphyU'U8; Zo, L'Upin'U8 ornat'U8; p, Poa 8andbergii;
e, Leptotamia m'Ultijida; a, Agropurum 8picat'Um.

8

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systems examined reached depths of over 10 feet; because of the devious
course they pursue, the tap is frequently much longer. Indeed, one was
found to have a length of nearly 13 feet. The abundant laterals, their wide
spread, and the clusters of terminal branches, together with the root nodules,
are shown in figure 7.
Lupinus leucophyllus.-This lupine has a much larger transpiring surface
but a much smaller absorbing surface than L. ornatu8, which it resembles in
having a well-developed tap-root, usually with several laterals, all of which
are branched to the third and fourth order. However, the root system is
much less extensive than that of L. ornatU8; its average depth is only about 5
feet (fig. 7).
Astragalus arrectus.-This legume forms marked estival societies, the
plants drying up during the first week in July. It has a strongly developed
tap-root, which usually pursues a course directly downward and sends out
many strong laterals at various depths. These laterals are fairly wide-
spreading and profusely branched to the tip. They reach depths of 4 or 5
feet (fig. 8).
Balsamorhiza balsam-root is a dominant in the high
prairies of eastern Washington. Its abundance, size, and duration all unite to
make it a very important species ecologically. From the short, thick, mul-
ticipital stem the new leaves appear in April. By the first of May the plant
is often in full bloom, forming societies which are very conspicuous, even after
the whole aerial part has dried up. Twenty-five root systems were excavated
and examined. Balsamorhiza has a tap-root sometimes reaching a diameter
of 4 inches and an extreme depth of 8 feet 10 inches. The laterals seldom
come off in the first 6 inches of soil, but numerous strong laterals occur below
this, sometimes 1 inch or more in diameter, and these often run rather hori-
zontally for 2 or 3 feet before they turn downward. They may ultimately
reach depths of 5 feet or more. The lateral branching is profuse, and in all
directions the soil is laid hold upon. Sometimes the tap splits up into nearly
equal parts at a depth of a few feet. The tip of the tap-root is often dead, and
if alive is never much branched. The older part of the root especially is
covered with a bark furrowed sometimes 0.5 inch deep. The average root
depth was found to be 5.5 feet (fig. 6).
Geranium viscosissimum.-This subdominant forms extensive societies in
the estival aspect. It has a well-developed tap-root which may reach 3 inches
in diameter. The tap sends off many laterals, both large and small, at all
levels, all of which-may branch profusely to the fifth order. The larger laterals
usually run off in a horizontal direction, sometimes for nearly 3 feet before
turning downward. The end of the tap-root is either unbranched or branched
but little, and is often dead. Hard soil seems to be a marked limiting factor
to root growth, and under this condition, especially, the usual very irregular
course of descent is greatly emphasized. Twenty-four root systems were
examined. While one plant penetrated to 9.5 feet, the average depth was
found to be 5.5 feet (fig. 6).
Wyethia amplexicaulis.-This large, rather coarse plant is at home on
moist hillsides and especially in valleys, where it forms extensive estival
sOcieties. It has a fleshy tap-root, which sometimes measures 9 inches in
circumference and may reach a depth of 6 feet 5 inches. It usually has several
strong laterals which come off from 8 inches to 3 feet in depth, and may run
out in a somewhat horizontal direction for 3 or 4 feet from the main root.
Often at a depth of 1 to 3 feet the whole tap breaks up into 2 to 5 nearly equal
"feet
3
2
1
2
3

FIG. 8.-Schematic bisect: hu, Heuchera glabella; a, A8tragal'U8 arrect'U8; 8, Sidalcea oreganai h, HeZiantheZla douglasii;
ag, Agropyrum spicatum.
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parts which pursue a downward course, or later extend out as laterals. Pri-
mary laterals are not much branched. The whole root from crown to near the
tip ,is covered with scattered small laterals, usually not over 2 rom. in diameter.
These are poorly branched, not often giving off roots of the fourth order. The
tip of the main root likewise is little branched. The roots dry out and shrink
rapidly upon removal from the soil. The 18 plants examined had an average
root depth of 5 feet (fig. 7).
Reuchera glabella.-This saxifrage has a strong tap-root supplied, especially
in the first foot of soil, with rather numerous but poorly branched laterals.
It reaches depths of about 5 feet (fig. 8).
Leptotrenia multifida.-Leptotamia is a subdominant which forms con-
spicuous societies in the vernal aspect. It has a large fleshy tap-root, some-
times 7 inches in circumference, which may reach a depth of over 5 feet. . The'
fusiform roots may narrow down gradually or rather abruptly, even to a
diameter of 1 or 2 mm., and then again enlarge to a size equaling the original.
This is sometimes repeated several times, thus giving the root as a whole a
beaded appearance. In general the roots are very poorly branched. The
average depth of penetration is 5 feet (fig. 7).
Relianthella douglasii-This species, like the following, forms societies in
the autumnal aspect. It has a tap-root with a diameter seldom more than 2
cm. It' throws out many large laterals just beneath the surface, most of
which come off within the first 18 inches of soil, although there are some
lower, and.a cone may be formed all the way to the tip. The laterals usually
pursue rather a vertically downward course and are profusely branched. Of
the 20 plants examined the deepest penetrated only to 5.5 feet, while the aver-
age depth was found to be 4.5 feet (fig. 8).
Roorebekia.racemosa.-The strong tap-root of this plant sometimes reaches
a depth of 10 or 11 feet, but usually ends near the 5.5 foot level. The larger
laterals, if any, are often thrown off within the first 18 inches of soil. These
are branched to the third order and may run off in a rather horizontal direc-
tion. The tap grows directly downward and is sparingly branched all the
way to the tip. The main root is usually not more than 0.5 inch in diameter
(fig. 6).
Lithospermum ruderale.-The tap-root of this species varies from 3 to 10
inches in circumference and gives promise of a deeper root system than is
actua.lly attained. Only one plant reached a depth of over 6 feet, while the
average depth of the 5 specimens examined was 4: feet 10 inches. Large
laterals were sent off from the main root at various depths from 1 to 2 feet.
These ran horizontally several feet before turning downward.
Sieversia. ciliata..-This species forms societies in the estivalaspect. Few
roots of Sieversia penetrate beyond depths of 5 feet 6 inches. It sends out as
many as 20 to 30 roots from a single inch of its thick rootstock. None of these
roots is over 3 to 4 mm. in diameter. They pursue a vertically downward
course and branch profusely all the way to the tip, sending off laterals seldom
over 3 inches long but branched to the fifth order. The 11 plants examined
gave an average root depth of 4 feet 9 inches (fig. 7).
Sida.lcea. oregana.-This mallow is confined to north hillsides and low
ground. It has a tap-root which gives off strong laterals, beginning in the
first foot of soil. The tap penetrates to depths of only 3 or 4: feet. The roots
are characterized by the origin of short laterals in groups of 3 to 8 (fig. 8).
ffieracium scouleri.-This hawkweed is one of the few species of the western
prairies which forms autumnal societies. For the 26 root systems examined
an average depth of 5.5 feet was determined. It sends out numerous roots,
as many as 50 from a single inch of its rhizome. The rhizome is 8 to 12 inches
long. None of these roots are over 3 mm. in diameter. They pursue a nearly
vertically downward course and throw off practically no laterals, except where
they enter earthworm holes. Here strong laterals, equaling the main root in
diameter, are developed and run parallel with it in the hole a foot or two or
further. All give rise to abundant root-hairs and it is seldom that these roots
again enter the soil. The tip of the main root may branch very profusely, or
very little if in hard soil. The longest root reached a depth of 7 feet 9 inches,
although 5 feet 4 inches was found to be the average depth (fig. 6).
Potentilla blaschkeana.-The tufted stems of this subdominant are borne
on a short, thick crown from which 2 to 5 main roots originate. These average
about 7 mm. in diameter. They taper off gradually till at about 2 feet in
depth they are often only 2 mm. in diameter. Here they usually branch
dichotomously, and again branching, break up into numerous small roots-
mere hairs-which have a strong tendency to grow in earthworm holes.
These they follow for perhaps 2 feet before entering the soil again. In these
holes they give off many threadlike branches which follow down the same
hole, branching profusely and often not reentering the soil. The roots are
dark brown in color. They are unbranched or very poorly branched at the tip.
Thirty root systems were examined. The longest roots penetrated to a depth
of 7.5 feet, while the average root depth was found to..be about 5 feet (fig. 6).
Eriogonum heracleoides.-This mat-fonner, at home on dry ridges, has a
strong, woody tap-root often an inch in diameter. It sometimes breaks up
just below the soil surface into several more or less equal parts. The roots
penetrate to a depth of 6 to 8 feet or more. They spread widely and are well
branched.
Some examinations were also made of the roots of R08a nutkana and Bym-
phoricarp08 racemo8U8, both very common in nearly all prairie situations.
They were found extending to depths of from 6 to over 8 feet. Iri8 mis-
80uriemis roots penetrated the soils of dry hillsides to 3 feet 10 inches, and
Berberi8 repen8 to a depth of over 10 feet.
PRAIRIE ROOT SYSTEMS AND PRAIRIE ENVIRONMENT.
From these data it may readily be seen that most of the prairie
plants are deep-rooted. With the exception of Kreleria cri8tata, Poa
8andbergii, and Fe8tuca ouina (which are confined largely to the surface
,18 inches of soil) all of the species examined reached depths of from 4
to 6 feet, while some penetrated even deeper. However, when com-
pared with the species of the eastern prairie, these plants are not as
deeply rooted. It may be recalled (p. 18) that about 55 per cent of
the roots in the latter community penetrated well beyond 5 feet and
many of them beyond 10 or 12 feet. Here, however, with a few
exceptions, the plants usually terminate their root systems at depths
of about 5 or 6 feet, while only one or two species have roots penetrating
to 10 feet. A proper interpretation of this difference can be made only
by a careful study of the habitat.
The prairies of southeastern Washington and their eastward exten-
sion into adjacent Idaho occupy a position between the foothills of the
Bitterroot Mountains on the east and the sagebrush region of western
Adams, eastern Franklin, and western Walla Walla counties of Wash-
ington on the west. On the south they are bounded by that high
upfold of the lava-rock known as the Blue Mountains. Northward the
Spokane gravels, extending somewhat southward of Spokane, with
their open growth of yellow pine, mark at the same time the general
northern boundary of the exposed part of the great lava sheet and its
accompanying prairie formation. Since the supply of water in this
region is the chief limiting factor to plant growth, we shall first con-
sider the total amount of precipitation, with its seasonal distribution,
after which the water-content of the soil will be considered.
Hemmed in on all sides by mountains, and especially cut off from
the moist winds of the Pacific by the Cascades, the Columbian Plateau
has a low annual precipitation. Even on its high eastern border,
where these studies were carried on, it is only 21 inches. Table 7
shows in inches the mean monthly and annual precipitation at Pull-
man, Washington.
TABLE 7.
Month.
Precipi-
Month.
Precipi-
tation. tation.
inche3. inche3.
Jan ...... 2.00 Aug .... 0.68
Feb ...... 2.18 Sept. .... 1.29
Mar ...... 2.02 Oct ... 1.70
Apr ..... 1.50 Nov ..... 3.41
May ... 1.84 Dec ...... 2.66
June ... 1.20
July .... 0.57 Total .. 21.60
It may be seen at a glance that about three-fourths of the precipita-
tion occurs during the resting period. The light showers of July and
August seldom have much influence on the water-content of the soil.
The soils of this region may be compared to a gigantic reservoir
replenished mostly during the resting season and rather thoroughly
emptied of its water during the summer. It is not the absolute rainfall
figures alone which furnish a criterion of climate, for the maximum
duration of the drought period constitutes a limiting factor of the
greatest importance. The great problem is the extent to which soil
water derived from the winter precipitation is conserved through the
weeks of drought. The rains in southeastern Washington are so gentle
that there is practically no run-off, and the silt-loam soils have a
wonderfully retentive power.
The prairie soil has originated from the decomposed underlying
basalt. It consists of a friable, dark-brown silt loam which has a water-
holding capacity of 50 or 60 per cent of its dry weight. By the action
of water and especially of the prevailing southwest wind, the prairie
topography has been molded into rounded hills which reach a height
of 100 to 360 feet and resemble sand-dunes. The soil is usually many
TABLE 8.
SW. slope. NE. slope. SW. slope. NE. slope.
Date. Wilting Wilting Date. Wilting Wilting
coefficient 11. coefficent 12.2. coefficient 11. coefficient 12.2.
1913. 1913.
Apr. 25 .. 10.0 21.0 Aug. 15 .... -4.2 0.0
May 2 .... 12.0 22.2 Aug. 22 .... -2.6 -1.3
May 8 .... 9.0 20.5 Aug. 28 .... -3.0 -1.0
May 14 .... 13.3 25.6 Sept. 5 .... -2.2 -2.0
May 20 .... 13.0 25.0 Sept. 10. '" -3.4 -1.8
May 27 .... 5.2 18.0 Sept. 17 .... -3.9 -1.0
June 2 .... 2.0 16.6 Sept. 25 .. -2.2 0.7
June 10 .... 4.8 21.2 1914.
June 15 .... 5.0 17.4 May 22. '" 4.0 13.0
June 25 .... 8.6 26.0 June 2 .... 0.1 7.5
July 2 .... 4.6 19.0 June 26 .... -1.3 1.3
July 8 .... 4.0 15.2 July 4 .... -0.2 1.8
July 16 .... -0.5 9.4 July
8. '"
-1.2 1.1
July 21. ... -0.5 6.4 July 19 .... -4.5 0.0
July 28 .... -2.6 4.4 July 27 .... -5.0 -2.0
Aug. 4 .... -1.4 0.2 Aug. 3 .... -4.0 -2.0
Aug. 9 .... -3.0 0.0 Aug. 10 ... ; -4.0 -2.8
TABLE 9.
SOUTEnWEST SLOPE.
Wilting coefficients at stated depths.
Date.
11.5 at 11.0 at 14.2 at 13.5 at 13.5 at 14.0 at
o to 6 n c h e ~ . 6 to 12 inches. 2 feet. 3 feet. 4 feet. 5 feet.
Dec. 13, 1913 ... ............ 11.5 -1.2 -0.3 0.5 0.5
Apr. 18, 1914 ... 14.5 13.5 8.1 9.7 8.2 5.5
June 3, 1914 ... -1.0 1.4 3.4 7.3 8.1 8.0
July 6, 1914 ... -2.-5 0.0 -0.7 1.0 5.1 6.8
Aug. 15, 1914 ... -4.5 -1.6 1.9 0.6 2.1 2.6
NORTHEAST SLOPE.
Wilting coefficients at stated depths.
Date.
12.2 at 12.2 at 13.4 at 13.6 at 14.2 at 14.0 at
o to 6 inches. 6 to 12 inches. 2 feet. 3 feet. 4 feet. 5 feet.
-
Dec. 6, 1913 .. ............ 15.8 11.2 -0.8 0.6 5.0
Apr. 18, 1914 ... 22.8 20.0 15.6 13.4 9.9 9.5
June 3, 1914 ... 6.6 8.5 13.1 16.5 13.6 9.8
July 6, 1914 ... 3.3 1.0 6.8 12.1 10.8 3.9
Aug. 15, 1914 ... -3.0 -1.7 -0.6 0.0 2.3 3.0
feet deep. The wind has drifted much surface soil and humus material
from the exposed south and southwest slopes and deposited it upon
the steeper north and northeast leeward slopes.
It may be seen in tables 8 and 9 that the soil in the spring shows a
maximum water-content, the autumn and winter precipitation having
replenished the water lost during the long period of summer drought.
Table 8 gives the available of the prairie soil to a depth
of 10 inches during 1913 and 1914. Table 9 shows the available water-
content of prairie soils to a depth of 5 feet during 1913-14. The minus
sign indicates water non-available for plant growth.
TABLE 10.
'May22. July 6. Aug. 15.
Depth.
NW. S. NW. S. NW. S.
----------
o to 6 inches ..... 23.4 12.5 15.2 11.1 11.6 6.7
6 to 12 inches ... 25.7 17.9 13.7 11.4 12.1 10.2
At 2 feet ........ 27.0 19.9 18.9 12.6 11.8 10.2
At 3 feet ........ 30.2 20.2 23.7 13.2 14.4 10.6
At 4 feet ........ 27.2 22.0 24.9 17.3 18.0 10.5
At 5 feet ........ 24.3 22.4 21.3 19.6 19.5 12.3
The greater water-content on northerly exposed slopes is the result
of several causes. More precipitation actually occurs here as far as
effective moisture is concerned, much of the 46 inches of normal snow-
fall being blown over to the sheltered north hillsides. The soil on
north slopes is deeper, has more humus, and a greater water-holding
capacity. It has been seen that this reflects itself both in plant dis-
tribution an,d root development. An examination of table 9 shows
not only a gradual depletion of soil moisture as the season progresses,
but also that to a depth of 2 or 3 feet little or no moisture is available
in late summer.
This seasonal march of soil-moisture, together with its greater
scarcity on south slopes, is further illustrated in table 10, which shows
the water-content of soil samples taken from another set of stations
during 1914.
To evade the drought conditions thus imposed upon them, a few
plants, notably the shallow-rooted grasses, complete their period of
growth and produce seed early in the season, and then lie dormant
until revived by the autumn rains. Most prairie species, however,
extend their roots far down into the deeper moist soil. As the moisture
supply decreases with the progress of summer, the evaporating power
of the air becomes more and more intensive. This is shown in table 11.
Thus, high evaporation and low available soil moisture combine
to make conditions rather unfavorable for plant growth, especially
in late summer and in exposed situations. As has already been men-
tioned, late-maturing species are not at all abundant. No late-
maturing grasses are components of the prairie flora. Flowering
begins later and growth continues much longer on north hillsides.
Soil temperatures on the two exposures differ notably. These data
are shown in table 12, which gives the soil temperature at 1 to 5 feet
on April 18 and August 15,1914, on a northeast and southwest slope
respectively.
TABLE n.-Average daily evaporation in cubic centimeters on a northea8t
and a southwest slope respectively.
Date. 8W.slope. NE.slope. Date. SW.slope. NE.s1ope.
1913. 1913.
May 10 .. 12.0 10.5 Sept. 3 .. 33.0 21.7
May 15 .. 9.8 6.5 Sept. 7 .. 32.1 21.6
May 26 .. 13.1 10.2 Sept. 13 .. 33.6 23.8
June 3 .. 15.2 10.2 Sept. 20 .. 46.0 20.0
June 10 .. 13.0 8.6 Sept. 23 .. 21.0 17.0
June 15 .. 14.2 7.8 1914-
June 25 7.8 5.4 May 15 .. 20.1 19.0
July 3 .. 11.0 6.7 May 21. . 19.0 15.1
July 8 . 21.0 13.5 May ao .. 17.8 15.6
July 13 27.2 14.0 June 7 .. 20.0 11.2
July 18 .. 31.0 21.8 June 15 .. 16.1 12.5
July 25 30.8 20.3 June 22 32.1 19.0
July 29 .. 41.5 26.4 June 29 .. 19.0 12.8
Aug. 4 . 33.0 23.8 July 6 .. 27.2 22.5
Aug. 10 .. 39.5 24.5 July 13 .. 44.0 30.5
Aug. 16 .. 26.5 15.8 July 20 .. 44.8 26.0
Aug. 20 .. 38.0 27.2 July 27 . 49.0 32.0
Aug. 25 . 44.3 30.0 Aug. 3 . 51.5 42.6
Aug. 31. . 43.8 27.5 Aug. 10 . 57.8 39.1
Aug. 15 .. 64.0 50.1
TABLE 12.
April 18. August 15.
Depth.
NE. SW. NE. SW.
1 foot .. 45. 7F. 50. 3F. 65.00F. 73.0F.
2 feet .. 45.1 50.0 61.7 70.0
3 feet .. 45.0 48.2 58.0 66.2
4 feet . 45.0 47.0 57.2 64.4
5 feet . 45.0 46.4 54.5 62.6
The daily midsummer range at a depth of 3 inches on the northeast
and southwest slopes was 12
0
F. and 26
0
F. respectively. It is probable
that these differences in temperature have considerable effect at least
on the rate of root development (Cannon, 1918).
Summarizing briefly the factors of the habitat in which ~ h e s e
studies were carried on, we find a . region of moderate winter and low
summer precipitation. The soils are composed of a fine silt loam of
high water-holding capacity and they are usually very deep. In early
summer the superficial layers of soil soon lose all of their water avail-
able for plant growth, and as the season advances this condition occurs
in the deeper soils, while the entire soil-mass (to a depth of 5 feet and
beyond) gradually yields most of its available water. Soil temperatures
at 3 inches show a daily range of from 3 F. to 24 F., while at 1 foot
the daily range is seldom over 1 F. The seasonal range (April to
August) of the soil temperatures varies from 22 F. at 1 foot to 16 F.
at 5 feet. Air temperatures show a mean daily range varying from
about 25 F. in April and May to 38 F. in July and August. The
cool nights on the high plateau tend to counteract the low humidity of
the day and to reduce the high daily rates of evaporation.
North and northeast slopes are less xerophytic than the south and
southwest slopes. This is due in part to actually greater precipita-
tion caused by blowing snow and in part to soil texture, which is more
open, has more humus, and a greater water-holding capacity. These
factors are reflected in the greater amount of soil-water and in lower
soil temperatures. Likewise these slopes are sheltered from the drying
southwest winds and from the perpendicular rays of the sun. This
is reflected in slightly lower air temperatures and greater humidity,
and especially in the lower evaporating power of the air. On the part
of the plants the most obvious response to these severe environmental
conditions is the development of extensive root systems. For just as
the evaporating power of the air and the nature of the transpiring
organs determine the water-loss of plants, likewise the soil-water
and the nature of the root systems determine the supply.
TABLE 13.-Average root depth8 (in inches) of plants on different slopes.
SW.s1ope. NE.s1ope. NW.slope.
Lupinus ornatus ....... 72.1 71.0 128.4
Lithospermum ruderale ... ........... 48.7 74.5
Potentilla blaschkeana ... .......... 59.2 64.3
Geranium viscosissimum .. .......... 56.1 72.2
Hoorebekia racemosa ..... 64.0 63.4 68.2
Hieracium scouleri ....... 62.8 63.4 66.8
Helianthella douglasii .... 52.0 55.6 55.1
Balsamorhiza sagittata ... 61.5 .......... 69.8
Leptotrenia multifida ...... .......... 54.7 47.3
Agropyrum spicatum ..... .......... 49.6 38.6
It is instructive to note that according to the types of root systems
as set forth by Cannon, all the roots here described, with the possible
exception of Leptotamia multifida, fall under the generalized class. This
was true also for all but a few of the eastern prairie species. By a
generalized root system is meant one that has botli the tap and the
laterals well developed. They penetrate deeply and reach out widely.
In contrast, the specialized root system has either the tap-root as the
chief feature or the laterals are placed near the surface and are espe-
'cially well developed, as in cacti. The generalized type of root is much
more plastic and consequently reacts to a wider range of conditions
than does the specialized type.
In considering the question of the susceptibility of roots to modi-
fication through variation in the soil texture or its water-content, as
against the conservative inherited tendencies, table 13 is instructive.
It appears that the root lengths on northeast and southwest slopes
are about the same and that the marked environmental differences
play little part in determining root depth. The greater root depth of
most species on the northwest slope at a station which combined a
porous moist soil with rather extreme xerophytic above-ground con-
ditions is at once evident. In all habitats studied the writer has found
that hard soil profoundly affects the amount of branching, laterals
practically always being more numerous in a less compact substratum.
This may be closely connecte.d with water-content and aeration-
the looser soils, of course, if'of the same type, having the greater water-
holding capacity. A discussion of the effects of various habitats on
the root development of polydemics will be found- on page 110.
The roots of several species showed a marked increase in their out-
put of branches upon leaving the compact soil and entering earth-
worm burrows. In practically all cases the root-tips under such con-
ditions were alive and at least well covered with root-hairs, while in
the more compact soils of dry slopes especially the root-tips were
often dead and decayed. The cause of these differences is yet to be
determined. The differences may be due to the mechanical resistance
offered by the soils, or to changed conditions of aeration, or perhaps
to both factors acting together. Cannon and Free (1917) have found
that roots of various plants respond quite differently to variations
in the composition of the atmosphere. This difference appears to be
related to the character of the natural habitat of the species in ques-
. tion. Their results indicate that plants growing in well-drained soil
are much more sensitive to the composition of the soil atmosphere
than those from poorly drained and poorly aerated habitats.
Contrasting grassland species of the east and west, we reach the
following conclusions: Prairie plants of eastern Nebraska growing in
the clay-loam or loess soil under an annual precipitation of 28 to 30
inches, most of which occurs during the growing season, and with an
average daily summer evaporation of 21 c.c., have root systems very
similar in distribution but somewhat greater in depth than prairie
species of southeastern Washington growing in the silt-loam of dis-
integrated basalt under an annual precipitation of 21 inches, most of
which falls during the resting season, arid with an average daily sum-
mer evaporation of 30 C.c. .
In both of these semiarid regions it has been shown that the water-
content of the soil is reduced to the non-available point to a depth of
3 or 4 feet, at least during 'certain years. It has also been shown that
in both of these prairie communities all of the dicotyledonous plants,
as well as many of the grasses, extend their root systems to depths of
from 3 to 7 feet or more.
The absence of very shallow, widely spreading laterals is charac-
teristic of plants of both prairie communities, perhaps being more
pronounced in the Palouse region. This is in distinct contrast to the
root habit of many plains species, to which we shall now turn our
attention.
46 THE' ECOLOGICAL RELATIONS OF ROOTS.
IV. mE PLAINS ASSOCIATION.
This great grassland community extends from northwestern Texas
and northern New Mexico to northern Colorado and Nebraska and
from the Rocky Mountains eastward to central Texas and Kansas,
wherE" it meets the prairie. While soil moisture seems to be the limit-
ing factor in the extension of the prairies westward, this is so inti-
mately connected with the amount and distribution of the precipita-
tion and with the soil type that great tongues of true prairie extend
far into the Great Plains. This is especially true northward, while over
much of the broad ecotone plains and prairie vegetation alternate.
The plains community differs from the prairie chiefly in the domi-
nance of short grasses, especially Bouteloua gracilis and Bulbilis dacty-
loides, and in the smaller number and reduced importance of the societies.
An excellent description of various plains communities, together
with their successional relations and indicator values, especially for
Colorado, has been given by Shantz (1911). Plate 18, A, affords a
glimpse of the plains vegetation near Colorado Springs, Colorado,
where the roots described below were excavated. Here about 80 to
85 per cent of the soil surface was covered with vegetation.
Bouteloua gracilis.-This well-known and important grass is a dominant'
over the plains from western Nebraska to the Rocky Mountains and from
Texas far into Canada. From the standpoint of grazing it ranks among the
highest of all the grasses, being equaled only by Bulbilis dactyloides.
The soil is well filled with fine rootlets to a depth of 30 inches, while in the
next 6 inches they are still fairly abundant, some of the longer ones penetrating
to a maximum depth of 48 inches. The surface roots spread very widely in
the shallow soil to 1.5 feet or more and are exceedingly well branched.
Aristida purpurea.-Much of the plains association is characterized by the
bunches of this very widely distributed dominant, wherever overgrazing or
other disturbance has given it a foothold in competition with Bouteloua gracilis.
From the bases of the individual clumps strong fibrous roots arise, from 0.5 to
1 mm. in diameter. These rather coarse roots either descend vertically or run
off obliquely at an angle of from 20 to more than 45 degrees with the surface
of the soil to a distance of 5 to 8 inches before turning downward. The sur-
face of the soil below the clump is completely filled with these cord-like roots,
.but they do not branch profusely until they have penetrated 4 or 5 inches into
the soil. Below this depth for 3 or 4 feet the roots become smaller, giving off
both large and small laterals, many of the latter being only 1 to 10 mm. in
length. The ground is rather well occupied to a depth of 3 feet, at which
depth many of the roots terminate in rather well-branched tips, while others
penetrate to a depth of 4 feet or slightly beyond. A maximum depth of 4 feet
3 inches was recorded for several of these tiny, hairlike termini. Branching is
especially well developed in joints between the lumps of hard soil. The older
roots are ashy-gray in color, and are provided with a rather papery cortex,
which upon removal reveals the tough, light-yellow stele. The deeper roots
are threadlike and are very fragile and usually well branched, although they
sometimes run for several inches without giving off any laterals (fig. 9).
Fissures occur in the soil, extending to a depth of 2 or 3 feet. These have
THE PLAINS ASSOCIATION
47
evidently been open cracks and the soil on the sides of them is much darker
in color, having been washed down from the surface. It is in these crevices,
where the moisture-content is somewhat higher, that grasses and other plants
branch and rebranch so profusely.
1 ft.--.j
FIG. 9.-Aristida purpurea. FIG. lO.-Artemi8ia frigida.
Muhlenbergia gmcillima.-This grass is also a dominant of the short-grass
plains; it is less widely spread and less abundant than Bouteloua gracilis,
though in some places it ranks close to the latter. Superficially, it appears like
Bulbilis dactyloides, because of its mat-like growth in small areas and the short,
curled leaves. Frequently the center of the mat is dead and only the
peripheral portions produce flowers.
A trench over 4 feet long was dug with one face cutting through a pure
stand of this grass to a depth of about 5 feet. Great clusters of roots, only
abput 0.5 rom. or less in diameter, ran off in all directions from the very sur-
FiG. ll.-Gutierrezia 8arothrm showing
wide lateral spread of roots in the
surface soil.
FIG. 12.-A small part of caudex of
Yucca glauca, showing number and
extent of horizontal roots, many
exceeding 25 to 30 feet in length.
THE PLAINS ASSOCIATION. 49
face to the average working depth, and are abundantly supplied with short,
much rebranched laterals. Many of these spread very widely laterally,
branching and rebranching into both long and short termini, so that the
surface 27 inches of soil is completely filled with a dense network of absorbing
. rootlets. Every cubic centimeter of the soil to a depth of 2 feet is occupied by
a network of this absorbing system. At a depth of 27 inches most of the root-
lets end in very much branched termini, but a few penetrate to greater dis-
tances. By undercutting the face of the trench and working upward from a
depth of 5 feet, several of these threadlike rootlets were encountered at a
depth of 55 inches. The roots are tan in color.
Gutierrezia sarothrm.-This half-shrub occurs throughout the plains region,
although it is usually more abundant in areas of less thoroughly disintegrated
soil or in overgrazed areas. In such situations it forms extensive societies.
The plants examined were quite abundant and from 6 to 9 inches high. Nu-
merous stems arise from a strong tap-root from 5 to 8 mm. in diameter. The
tap immediately gives rise to 5 to 9 large laterals, 2 or 3 mm. in diameter, and
as a result, at a depth of 2 to 6 inches the tap is no larger than one of these
laterals. The main root descends more or less vertically and at a foot in depth
is seldom over 1 mm. in diameter. It is much curved and kinked, but these
twists are usually small, so that at a maximum depth of 6 or 6.5 feet the tip
of the root is not more than a foot or 18 inches horizoritally from the base of
the stem. Just at the surface, to a depth of 12 to 18 inches, very numerous
threadlike laterals, 0.5 mm. or less in diameter, run off horizontally in all
directions for a distance of 3 to 24 inches, the plant thus being well supplied
with surface feeders. The large laterals frequently pursue a course nearly
parallel with the surface of the soil at a depth of 2 to 5 inches, or run some-
what obliquely to distances of 12 to 20 inches before turning rather abruptly
and vertically downward. Like the tap, they are profusely branched and
rebranched with short, threadlike laterals, but after a depth of 18 inches is
reached the branching becomes much poorer. These deeper roots may run a
distance of a foot or more without giving off any laterals, or again short laterals
.only 2 or 3 inches long may occur in groups. The tips of these deeper roots
are only fairly well branched. Some were found which gave off a whole net-
work of branches matted in the cracks. While the larger roots are brown,
tough, and rather woody, the deeper ones are nearly white and rather brittle.
Several roots were traced to a depth of over 5 feet, while a few reached a
maximum depth of 6 to 6.5 feet (fig. 11).
Psoralea tenuifiora.-This is one of the important legumes throughout the
plains. Under favorable conditions of soil moisture it forms extensive societies.
A number of individuals of this species were examined. They all had strong
tap-roots varying in diameter with the age of the plant. One large root,
which unfortunately was dead, had a diameter of 2 cm. At a depth of 1.5 feet
it gave off numerous large lateral branches, the branching system being not
unlike that of the more robust form of the prairies. A younger specimen,
with a tap-root 7 mm. in diameter, pursued an almost vertically downward
course to a depth of 42 inches. At a depth of 2 feet a small lateral ran off
rather horizontally to a distance of 20 inches. No other branching occurred
except near the tip, where 5 large branches and several smaller ones originated.
Except for a very few rootlets, 5 to 10 mm.long and distributed very irregularly
along the course of the tap, no other branching occurred.
A large plant, with a tap 8 mm. in diameter, reached a depth of 8 feet.
Like the former, it pursued an almost vertically downward course. It was
unbranched to a depth of 28 inches, where it gave off a horizontal lateral only
1.5 mm. in diameter and 27 inches long. A similar lateral about the same
size aroSe 6 inches deeper in the soil. At 5 feet the tap, now only 1.5 mm. in
diameter, forked dichotomously. At 6 feet the two branches entered a layer
of moist sand through which they ran for 26 inches to a total depth of 8 feet,
but they gave off very few branches. Rootlets of other plants-of this species
were quite numerous in these deeper soils. Finally, a plant was excavated
and photographed (plate 18, B) which reached a depth of 12 feet 2 inches.
The roots are brown to reddish-brown in color.
Artemisia frigida.-Mountain sage is a competitor of the plains grasses,
especially northward, for the rather meager water-supply. Near the moun-
tains; in rocky or gravelly situations, it frequently forms extensive societies.
From the base of the clustered woody stems a tap-root arises; it is from 5
.to about 10 mm.. in diameter and descends rather vertically to a distance of
froin 4 to over 6 feet. The tap, however, becomes rapidly attenuated, so that
at a depth of 6 inches it is scarcely 1 to 2 mm. in diameter and indeed no
larger than the . numerous strong, rather horizontal laterals which arise from
the tap or the base of the woody stem at just below the ground surface. These
larger laterals, frequently as many as 5 to 9 on a single plant, run off obliquely
in the shallow soil, sometimes almost horizontally and from a depth of only
1 to 2' inches to a distance of 8 to 12 inches or more before turning rather
abruptly downward. The woody bases of the stem, as well as the first 6 to 10
inches of tap-root, give rise to very abundant rootlets, 1 tmn. or less in diameter,
which thoroughly fill the surface soil, branching and rebranching into thread-
like termini and furnishing 'Artemisia, as is the case with so many plains plants,
with a splendid surface absorbing system. .
Like the tap-root, the larger laterals upon turning downward become
greatly attenuated, the diameter remaining uniformly a millimeter or less for
several feet. When near the ends they often becoJIle hairlike but are not well
branched. The deeper roots, while branching from time to time and pursuing
a rather tortuous course through the soil, are characterized by the absence of
numerous laterals. A large number of plants were examined and most of the
major branches extended with the tap to depths of from 3 to 6 feet. The
deepest root examined entered at a depth of 5 feet into the soft soil filling an
ancienthurrow, in which it continued, giving rise from time to time to long,-
threadlike, unbranched laterals. It reached a depth of 7 feet 9 inches. The
roots are chestnut-brown in color and all but the larger ones are very brittle,
being removed with extreme difficulty from the hard, lumpy soil (fig. 10).
Argemone platyceras.-This wild poppy, conspicuous because of its targe
size and showy white flowers, is found as a frequent component of plains
vegetation throughout much of the association. As a rule, it is more abundant
in disturbed areas, often becoming ruderal.
. The plant examined had a strong tap-root. It was partially decayed, por-
tions of it being dead and easily dug out. At the end of the first foot, however,
it branched dichotomously. These branches diverged not more than 6 to 8
inches and took a downward course, continuing to branch dichotomously at
irregular intervals for the first 8 feet, the whole trend being downward (fig.
13). At no point in the first 8 feet of soil did the branches spread more than
1.5 feet from the vertical. At about 8 feet a layer of moist sand was encoun-
tered and here some of the branches turned outward almost horizontally,
extending 18 to 24 inches, where they ended. Other branches continued
downward through moist sand to a maximum depth of over 12 feet, the last
10 to 12 inches of their length passing into a very moist sandy clay. From
THE l'LA3S ASSOCIATION. 51
about 4 feet downward these branches gave off occasional laterals about 2 to
4 mm. in diameter, which extended from a few inches to a foot or more. These
laterals tended to take a more or less horizontal direction.
The color of the root is dark brown. It was very thick and fleshy and
extremely brittle, so much so that near the terminus great difficulty was
. found in following it. Throughout its length, whenever broken, a clear to
milky fluid exuded in limited quantities. The surface of the root was extremely
rough and pitted; its diameter was often irregular. Throughout the whole
course.l\Vo or more branches bad a tendency to twist about each other. This
was marked in branches occurring not more than 2 feet from the surface,
again at intermediate depths, and stiIl again at not less than 6 inches from the
extreme depth.
Yucca glauca.-The soap-weed is a widely distributed species, being
especially abundant on dry, sandy, rocky slopes and forming extensive
societies throughout many areas in the plains association. Its size, duration,
and often its abundance combine to make it an important species. From the
standpoint of the stockman it is often a bad weed.
A number of specimens were examined. An excavation was made about
two large plants growing 2 feet apart. They each sent down a strong caudex
3 inches in diameter, but neither of them reached a depth greater than 18
inches. At this depth they branched and ran off laterally in a direction either
parallel with or ascending toward the surface. The two plants were connected
by a large underground rootstock (plate 17, A), while another (over 2 inches
in diameter) ran off at nearly right angles from the connecting rootstock and
reached the surface 3 feet beyond. It had given rise to several small plantlets.
From the multicipital stem as many as 25 to more than 50 rosettes of leaves
arose. Many of these were dead. One crown measured 21 by 26 inches in
diameter, this being only a medium-sized plant. It reached a height of 30
inches. The multicipital branching extended to a depth of 5 to 7 inches an,d
profound contractions were in evidence.
Beginning at the very surface, the stems are supplied profusely with roots.
These vary from 2 to 4 mm. in diameter and run off horizontally to great
distances, the lateral spread being remarkable. A number of these hori-
zontal roots were traced to a distance of 32 feet (fig. 12). They occupied the
soil chiefly at a depth of 6 to 18 inches, and, because of a somewhat tortuous
course, terminated at a horizontal distance of 27 feet from the caudex from
which they arose. These roots are reddish in color, of nearly uniform diameter
for long distances, and very sparingly branched. Only at intervals do short,
usually unbranched, secondary laterals occur. The cortex is thick and fleshy
and serves for water storage; upon the death of the older roots it shreds off
and reveals the tough wiry stele. The abundance of these roots is surprising.
In a single square foot of vertical trench face, at a depth of 18 inches, 54 roots
were counted. They extend outward in all directions from the rhizome, as
was proven by digging a series of small trenches at distances of 18 to 20 feet
on all sides of isolated specimens. Invariably Yucca roots were found. Such
a trench, dug halfway between two isolated Yucca plants 66 feet apart,
revealed the presence of laterals. Thus it may be seen that these large plants
absorb water and soil solutes over a very extensive area in direct competition
with the grasses.
A trench 6 feet long and 7 feet deep revealed very few Yucca roots.below 2
feet. However, some do occur and penetrate to maximum depths of about
7 feet. Like the shallower laterals, the deeper ones are poor1.ybran:cnea,even -
~ i } : > s having few branches and ending abruptly. The ends are not more \
than 0.25 to 0.2 mm. in diameter. At a depth of 4 or 5 feet, a few roots were i
found running horizontally, simulating those of the shallower soils. '
Another plant had a caudex 3 inches in diameter which reached a depth of
23 inches. Here it was still 2 inches thick. It branched into two equal
laterals. These were much flattened and distorted, one being less than an
inch thick but 2 inches wide. These branched and rebranched, as shown in
plate 18, c, all ending within a horizontal distance of 16 inches from the base
of the plant. Each was abruptly tipped with a bud. These deeper branches
were not so well provided with roots as was the upper part of the stem.
Agropyr8ln grass, if at all abundant, is an indicator of
favorable deep soil-moisture conditions. However, in thin stands and as
dwarfed individuals it occurs in rather dry places. The plants here described
were growing in low ground near a ravine.
The tufts of these coarse plants are connected by stout, tough rhizomes
about 2 mm. in diameter and from a few inches to more than 18 inches in
length. From the base of these clumps and from the rhizomes, which lie at a
depth of about 1 inch, arise numerous short horizontal roots. These are pro-
fusely branched and rebranched to the third and fourth orders, the ultimate
branches being almost microscopic in size, and thus furnishing a splendid .
surface absorbing system.
An abundance of coarse, tough roots, from 1.5 to 2 mm. in diameter, pene-
trated in a more or less vertical direction and others at an oblique angle to a
maximum depth of over 7 feet. These coarse roots are covered with a brown
papery cortex, rather readily removed from the 1arge w:h:e-like nearly white
stele. These roots are profusely branched with laterals from a few milli-
meters to 2 or 3 inches long, many of these branches running off more or less
horizontally. At a depth of about 4 feet many of these roots become only 0.2
mm. in diameter, but the branches (which are somewhat shorter) are no less
abundant to the very tip. The roots have such an abundance of root hairs
that the whole surface appears to be covered with wool. Thus Agropyrum
is supplied with an absorbing system which thoroughly permeates all portions
of the soil to a depth of 6 feet.
Carex pennsylvanica.-This sedge has a wide range throughout the grass-
land formation. Because of its early growth and flowering habit, it forms
conspicuous societies in the prevernal aspect, often before the taller grasses
reswne growth.
The roots of a number of plants of this species were examined. The tufts
are connected by coarse rhizomes, 2 to 10 inches long, at a depth of 1 to 3
inches. The much-branched fibrous roots have a lateral spread from the base
of a tuft of only 2 to 3 inches. They originate from the rhizomes as well
from the base of the clumpil. While many of the roots, after sending off
abundant laterals which branch to the third and fourth order, end at a depth
of 12 to 14 inches in exceedingly well-branched tips; others penetrate deeper.'
A few roots were traced to their delicate endings at a depth of 3 feet, while
others occurred below the second foot. The chief absorbing area lies within
the first and second feet of soil.
Andropogon seoparius.-The little bluestem occurs throughout the grasS:
land formation wherever enough water is present to support its growth. In
the drier plains association it is often confined to slopes with a rough topog-
raphy and greater water penetration or to sandy soil. It dominates the short-
graSses under the latter soil condition and forms an extensive bunchgrass
consocies. Under the still more favorable moisture-supply of the prairj.es it
abandons the bunch habit and becomes a sod-former. The lateral spread of
the roots is about 1 foot on each side of the plant and within 4 inches of the
surface. The roots are very abundant up to the very surface, thus affording a
THE PLAINS ASSOCIATION.
53
distinctly shallow absorbing system. Many of the laterals run out to a dis-
tance of 12 to 14 inches and at a depth of only 1 to 4 inches, while below and
inside of these at all angles to the vertical they are very abundant. The
maximum depth for several roots was 6 feet. They are very abundant to a
depth of 3.5 feet. The surface absorbing roots were especially well branched.
Stipa coma.ta..-This grass often appears to be a dominant in the plains
association where overgrazing has not thrown the advantage in favor of the
short grasses. Although its aerial part is not extensive, it has a much deeper
and better developed root system than the corresponding species of the
prairies, S. spartea. A trench 6 feet long and 5.5 feet deep was dug in a nearly
pure Stipa community. From the base of the clumps arise exceedingly
numerous fibrous roots, 1 mm. or less in diameter. Whlle many of these
desCend vertically, others run off at various oblique angles, some of them
having a lateral spread of more than 18 inches from the base of the plant,
while at a distance of 14 inches horizontally .they reach a depth of 6 inches
(plate 19, A). Beginning at the very surface of the soil, the main roots are
clothed with relatively short but well-branched laterals, 2 to 20 mm. long.
Thus the first 28 inches of soil are thoroughly occupied by the main roots,
which are only a few millimeters apart, the interstices being completely
occupied by horizontal branches. Even to a depth of 32 inches the soil is
fairly well filled with much kinked and rebranched threadlike rootlets, while
not a few reach a maximum depth of over 5 feet. Thus Stipa comata is pro-
vided with a much finer, more branched, more widely spreading, and deeper
. root system than is Stipa spartea of the prairies.
Lithospermum linea.rifolinm.-This is a common and rather' conspicuous
plant of the grassland formation. Eastward it forms vernal societies. The
plant examined had a crown of 10 stems arWng from the top of the tap-root,
which was 13 mm. in diameter. The tap proceeded straight downward 2.5
inches, where it divided into two equal brancheS, each about 7 mm. in diam-
eter. These' branches then took a downward course, their diameter decreasing
very rapidly, so that at a distance of 22 inches from the surface they were not
more than 1.5 mm. wide. This diameter remained almost uniform for several
feet. They continued their downward- course almost vertically, one part
branching again at a distance of 3.5 feet and the other at a depth of 5 feet.
These .branches of the second order aha took a downward course, being not
more than 0.8 mm. in diameter. Lower down, one of these branches again
branched and penetrated to the maximum depth of 10 feet, ending in hairlike
termini. The root was very dark, almost black in color, and had a papery
cortex which could be removed very easily, revealing a white hard and very
brittle stele. Throughout the lower 5.5 feet the roots were fairly well supplied
with short, attenuated branches 0.5 to 3 cm. long. Above this point shorter
branches were few in number. Very little absorption took place in the first
4 feet of soil (fig. 14).
Lygodesmia juncea.-Although this plant is of frequent occurrence, espe-
cially in drier situations, it plays a r6le of no great importance in the grass-
land. It is interesting because of its xeroid-shoot habit. Two plants were
examined. Each had a tap-root about 7mm. in diameter, which took an
almost vertically downward course, in one plant penetrating to a depth of
5 feet 8 inches. At 10 to 12 inches from the surface strong laterals were given
off. which (after taking a horizontal course from 8 to 22 inches) turned down-
ward not unlike the tap-root. Practically no small branches were given off
and only very seldom, as shOwn in figure 15, did any branching at all occur.
The roots throughout, both tap and laterals, pursued a very zigzag and tortu-
'\
> I
~
FIG. 13.-Argemone platyceras.
FIG. 14.-Lithospermum
lineari/olium.
ous course. The roots were dark brown with a very fleshy cortex, extremely
brittle, and when broken exuded a milky latex. At the top the diameter was
about 5 mID. As the root proceeded downward the diameter fluctuated from
4 to 7 mm., but gradually grew f\maller to the tips, which were about 1 mm .
.in diameter and ended abruptly. These roots are very different from the
much longer but almost unbranched roots of the same species growing in
loess soil (p. 16).
Aragallus lambertii.-This loco unfortunately is a widely distributed legume
throughout the plains grassland, forming one of the most important of the
vernal societies. As is characteristic of most legumes, AragaUus has a strong
tap-root. At the top this was 9 mm. in diameter and with a long gradual
taper it penetrated the soil with a tortuous course almost vertically downward
to a maximum depth of 8 feet. For the first 4 inches of its course there were
scarcely any branches except 3 or 4 small hairlike laterals, which penetrated
horizontally for a distance of 2 or 3 inches. Below this point and to a depth
of 30 inches the tap was marked by rather profuse, small laterals, ranging
from hairlike to 0.7 mm. in diameter. These penetrated the earth in a more
or less horizontal direction for a distance of 2 to 8 inches; they were them-
selves branched to the second or third order. Below 30 inches the tap was
very sparingly branched throughout its course, these branches being of the
same character but much smaller than those already described. The tap
ended by dividing into 3 or 4 ultimate rootlets, 3 or 4 inches long. The root
was tough, rather woody, light brown in color. Three plants were excavated
and examined (fig. 16).
Petalostemon purpureus.-This and the following subdominant have a
wide distribution throughout the grassland formation. Their presence in the
plains association is indicative of at least fairly good soil-moisture conditions.
They reach their best development in the prairies, where they form typical
societies. . .
More than a dozen plants were examined. They have tap-roots varying
from 3 or 4 mm. to 1 cm. in diameter. The general root system of all was
very similar. At a depth of 1 to 4 inches they invariably throw off 3 to 7
large laterals in a rather horizontal direction. These run off in the shallow
soil almost parallel with the surface. and often not more than an inch below
it, to a distance of 8 to 18 inches before turning rather abruptly downward.
Usually 3 or 4 of these laterals are more pronounced than the others, some of
them being as large as 6 mm. in diameter. Like the tap-root, they taper very
rapidly and throughout their horizontal course give off both large and small
sublaterals. These are frequently minutely branched and compete with the
grasses for the water in the surface soil. Upon turning downward, these
laterals, now usually not more than 1 to 2 mm. in diameter, pursue a course
in general vertically downward, though more or less curved backward and
forward. The tap-root descends vertically or runs off 6 to 12 inches from
this direction like the laterals, and tapers so rapidly that at a depth of 1 foot
it is seldom more than 1 or 2 mm. in diameter. Below this level, all the roots
are more or less threadlike, branching profusely, especially with small
rebranched feeders varying in length from a few millimeters to several inches.
The depth of the lateral branches usually does not exceed 4 or 5 feet, although
some were traced to a depth of more than 6 feet. The maximum depth of a
tap-root 6 mm. in diameter was 5.5 feet, while another 9 mm. in diameter
reached a depth of 6.5 feet. Unlike the white prairie-clover, this plant is well
adapted to absorb the moisture in the surface soil, but like the former the roots
occur quite abundantly to a depth of 5 or 6 feet, through a cross-section of 3 or
FIG. 16.-Aragallm lamoortii.
4 feet square under the plant. They are orange-brown in color, the deep toots
especially being easily followed and distinguished from their competitors by
this character (fig. 17).
Petalostemon candidus.-A group of more than 8 stems, all in blossom,
arose from a woody tap-root 17 mm. in diameter. At a depth of 3 inches this
tap split into three strong parts, 5, 6, and 8 mm. in diameter, respectively. A
larger branch ran obliquely for a distance of 3 inches and divided into two
equal parts, each 3 mm. in diameter. One of these descended rather verti-
cally, but with long, loose loops, 2 to 4 inches in width, turning back and
forth as was characteristic of other roots examined. The second fork ran off
obliquely about 4 inches further before turning downward. At a depth of 12
to 18 inches both branches divided and rebranched again and again, so that
at the 1S-inch level none of the roots were more than 1.5 mm. in diameter.
Below the second foot the roots ran for long distances, following much curved
and tortuous courses but with little change in diameter. The branches that
were given off at frequent intervals were very long and not much branched.
Finally, in the soil from the third to the fifth foot the branches became more
numerous and almost microscopic, the root-tipo being well supplied with
laterals only a few millimeters in length.
The other two main branches were very similar to those described, and
spread in such a manner that a cross-section of the absorbing area under the
plant would not include more than 4 square feet. Several of the hairlike
ultimate rootlets reached a depth of over 5 feet; the deepest one examined,
that of the vertically descending tap, reached a maximum depth of over 5.5
feet. After examining about a dozen species of each color, it was found that
the purple prairie-clover had many more superficial branches, which lie nearer
the surface. The roots are yellower in color, the older ones being almost
black. The branches are fewer, seldom more than 3, and run downward at a
much sharper angle. The types described are very characteristic. Thus it
may be seen that the white prairie-clover, Petalostemon candidus, in com-
parison with the purple prairie-clover, Petalostemon purpureUB, is not supplied
with absorbing roots in the surface 18 inches of soil, but gets the bulk of its
water and nutrients below this depth. The chief difficulty encountered in
excavating these plants was that of following the minute, dark-colored
termini in the third to the sixth foot of soil (plate 19, B).
Eriogonum jamesii.-This species is widely distributed over the plains,
where it forms summer societies. The thick woody root, 3 em. in diameter,
gave rise to a large number of prostrate stems which formed a mat about 8
inches square. Within the first 6 inches of soil, 10 Ot 12 laterals, the largest
of which was 5 mm. in diameter, ran off horizontally for a distance varying
from a few inches to 2 or 3 feet before turning downward. The tap tapered
gradually, so that at a depth of about 3 feet it was still 5 mm. in diameter.
To this depth it also gave off 2 other strong laterals and numerous smaller
ones, as in the surface 6 inches of soil. The former pursued a sinuous course,
such as is characteristic of Eriogonum roots, to a depth of several feet. At a
depth of about 3 feet, 2 other laterals arose which were only slightly smaller
than the tap. With the tap and rarely more than 12 to 18 inches from it,
these pursued an irregularly downward course, often turning backward and
forward horizontally or almost so, through a distance of 6 to 8 inches or more.
In the main, however, these roots as well as the tap had a vertically descending
direction. They were very poor1y branched, rarely giving off small unbranched
wirelike laterals. The roots were traced to a depth of 7 feet 3 inches, where
they were still 2 or 3 mm. in diameter, respectively, and they undoubtedly
58
,
penetrated several feet further. In color the o o ~ is red?ish brown and covered
with a scaly, shreddy cortex, the older parts bemg qwte woody (plate 19, c).
Ratibida columnaris.-This species is widely. distributed throughout the
grassland formation, where it forms summer societies dominating large areas.
It has a strong tap-root, 8 to 10 mm. in diameter. For the first 5 inches it
tapers very rapidly until at the end of
that distance it is not more than 3 mm.
in diameter. From that point it passes
vertically downward with a zigzag,
irregular course, very crooked and
crinkled, tapering slightly (at 12inches
being 1.5 mm. in diameter) to a max-
imum depth of 2 feet. The first 4 or 5
inches is marked by an extraordinary
number of more or less horizontal
branches which vary in diameter from
0.2 to 0.5 mm.; these extend out
through the surface soil to a distance
of 6 to 12 inches and are themselves
profusely branched with small hairlike
rootlets from 0.5 to 2 cm. long. The
number of these primary horizontal
branches is very large; one count
showed approximately 13, while
another showed about 40. Below 4 or
5 inches the root is seldom branched,
FIG. 18.-Ratibida columnaris.
what branches there are being hairlike and from 1 to 3 em. long. Near the
end, however, the tap usually branches into two or three small, short rami-
fying parts. The root is dark brown in color, rather tough and unyielding.
Four plants were examined (fig. 18).
Senecio aureus oblanceolatUB.-Senecio is a very common and often abun-
dant plains species, frequently forming vernal societies in the graPsland. The
specimens examined were small plants 6 to 10 inches high. The multiple roots
are so finely divided as to be almost fibrous. The number originating from
the crown varies from 15 ol" less to 30 or more. Some of these roots take a
downward course from the crown, but many of them pass out at a slight angle
from the surface to a distance of 4 to 12 inches. The point from which they
start variel'! from a few millimeters to 2 inches below the surface of the ground,
and the point at which they turn downward varies from 2 to 8 inches. These
roots then penetrate to a maximum depth of over 3 feet. The diameter of
these roots at the starting point varies from 0.2 to 1 mm. Throughout the
surface soil they are branched and rebranched to the third and fourth orders,
and each group of branchefl varies from 1 mm. to 3 inches in length. Below
6 inches these main roots, after starting downward, taper until they are not
more than 0.2 mm. in diameter. They are more or less branched, the branches
being alniost hairlike in diameter and 3 to 10 mm. long. In color, the roots
vary from a very light brown to a pure white. They are extremely brittle and
easily broken. Five plants were examined (fig. 19).
Asclepias verticillata pumila.-This dwarf milkweed is generally distributed
throughout the hard lands of the plains. It forms extensive midsummer clans.
The stems vary from 3 to 5 inches in height and are connected by an extensive
underground system which varies in diameter from 1 to 2 mm. These con-
60
necting stems extend from 2 to 5 inches in depth and range in distance between
the plants from 1 to 15 inches or more. At the points from which the above-
ground stems take their origin, one or two roots were found which penetrated
the soil, usually vertically but occasionally obliquely downward, following a
tortuous course. These vertical roots at the outset may range from 0.5 to
2 rom. in diameter and gradually taper downward to their extremity, which is
more or less branched and hairlike. The maximum penetration found was 46
FIG. 19.-8enecio aUTeus oblanceolatua.
inches. All parts of the root system, beginning about 2 inches below the soil
and extending to a depth of about 10 inches, are thickly supplied with small
laterals, usually about 0.3 mm. in diameter and from 0.5 to 3 inches in
length. These small laterals often come off in groups of 2 or 3. They are
themselves branched, ending in very minute capillary termini, thus affording
a very excellent system for surface absorption. The roots are white in color,
herbaceous, and for roots of this character fairly tough. When cut, small
amounts of latex exude. Six plants were examined (fig. 20).
Opuntia ca.manchica..-This Opuntia enjoys a very wide distribution in the
plains grassland. Its water requirement seems to be somewhat less than that
of Bouteloua. It is favored in its competition with the grasses by grazing, and
its development in great abundance is frequently indicative of overgrazing.
A single, carefully selected plant of average size was examined. As is
characteristic of the cacti in general, the root system consists of two distinct
parts-a few vertically descending anchorage and deep absorptive roots, and
FIG. 2O.-Underground parts of Asclepias verticillata pumila.
a much more extensive and shallow surface absorbing system. An examina-
tion of figure 22 shows 23 roots which run off in the surface soil, usually at
a depth of about an inch and seldom deeper than 3 inches, to distances
varying from 6 inches to 6 feet. The two largest roots were 5 mm. in diameter
and tapered very slowly in spite of the repeated branching. They were
branched repeatedly from their origin at the base of the plant to their
extremity with both large and small branches, which ramified in all directions
and thus furnished an enormous absorbing surface. The ultimate root end-
ings, whether of the shorter or larger branches, consisted of much-branched
and very delicate brushlike termini, to which the soil clung with great tenacity.
The smaller branches, except for their lesser extent, are similar to those already
described.
The deep anchorage and absorbing system of this plant consisted of 4 main
roots, which at the outset were 2.5 to 3 mm. in diameter. These extended
almost vertically downward, following a more or less irregular course through
the hard soil, branching as shown in figure 21 and reaching a maximum depth
of 35 inches. While the shallower roots were brownish in color and very
tough, these deeper roots were glistening white and much more fragile (cf.
Preston, 19(0). .
FIG. 21.-opuntia camanchica, FIG. 22.-Top view of surface roots of Opuntia caman-
showing vertical anchorage chica, showing the wide lateral spread.
roots.
Opuntia fragilis.-This cactus has a distinctively shallow root system. On
all sides roots extended out from the base of the plant, ranging from 1 to 2 mm.
in diameter, some having a lateral spread of 16 inches. There were about 20'
on the plant examined. Most of these took a course parallel with the
surface of the soil and lay within 3 inches of the surface, but a few took a
slightly more downward course, penetrating to a depth of not more than 8
inches. The first 3 or 4 inches of the root were sparingly branched, the few
branches being mostly mere tufts of rootlets not more than 3 or 4 mm. in
length. Throughout the rest of their course these divergent roots were
abundantly supplied with a system of laterals 0.2 to 0.3 mm. in diameter and
1 to 12 cm. long. These laterals were branched and rebranched to form
exten!?ive tufts interspersed throughout the soil, the whole forming a most
excellent system for surface absorption. Another plant gave a root depth of
15 inches.
PLAINS ROOT SYSTEMS AND THE PLAINS ENVIRONMENT.
An examination of the foregoing data impresses one with the great
depth to which the roots of most plains plants penetrate. There are
only 3 species of shallow-rooted plants that seldom extend below the
first 2 feet of soil (Kreleria cristata, Opuntia fragilis, and Ratibida
columnaris). Perhaps Opuntia camanchica should also be placed here,
though its vertically descending roots reach depths of 3 feet. This
group constitutes only 11 per cent of the 28 plains species examined,
while in the prairies 6 grasses or 18 per cent of the 33 species ex-
cavated had shallow roots. In the second group, made up of plants
with roots extending well below the second foot of soil but seldom
deeper than 5 feet, are found 32 per cent of the plains species as against
27 per cent among prairie plants. The third group of deep-rooted
plants, which extend to depths greater than 5 feet, contains 57 per
cent of the plains plants and 55 per cent of prairie species.
On the other hand, certain prairie plants, such as species of Rosa,
Lygodesmia, Liatris, and Kuhnia, reached depths of 17 to 22 feet. This
was quite beyond the greatest depth (about 13 feet) attained by any
plains plant. It may be that the soil is never moist to greater depths on
the plains. Moreover, most of the plains species are characterized, in
addition to their great depth, by a fine system of surface absorbing
and wide-spreading laterals. However, we must again resort to an
examination of the habitat factors in order to properly interpret the
root development.
The region in which these plants grow has an annual precipitation
of only about 15 inches, an amount quite typical for much of the
Great Plains area. Fortunately, the major portion falls during the
growing season. There is as a rule considerable rainfall during the
months from May to September, but often the rainy season covers
only June, July, and August. The mean monthly and annual precip-
itation in inches for Colorado Springs, together with its seasonal dis-
tribution, is shown in table 14.
Such a seasonal distribution of moisture is very favorable for the
growth of grasses. Because of the great compactness of the soil, brought
about in part by the binding influence of the short-grass cover, the run-
off is very high. Shantz has shown that at Akron, Colorado, the run-off
varies from 15 to 43 per cent, the greater run-off naturally occurring
during heavier rains. This combination of a meager amount of
precipitation with a relatively low degree of penetration would lead
one to think the soil must be quite dry. Weekly soil-moisture deter-
minations at various depths for the summer of 1918 are shown in table
15, which gives the available water-content during the summer of
1918. The minus sign indicates that no moisture was available for
plant growth.
TABLE 14.
Time. Amount. Time. Amount.
Jan. .... 0.20 Aug .. 2.20
Feb ...... 0.32 Sept ...... 1.17
Mar ..... 0.75 Oct. ...... 0.67
Apr ...... 1.55 Nov ...... 0.35
May ..... 2.43 Dec ...... 0.25
June. .. 1.88
July ..... 2.81 Annual .. 14.58
An exs.mjD8.tion of these data shows that, aside from variations in
the surface foot, the soil to a depth of 7 feet was rather uniformly dry .
How much' more deeply the rain may penetrate during a series of
wet years was not determined. Certainly, available moisture must
occur at least locally to depths reached by living roots of Psoralea,
Argemone, and other species which penetrate to 10 or 12 feet. Only a
Date.
June 7 ...... .
June 14 ...... .
June 26 ...... .
July 1 ..... ..
July 8 ..... ..
July 15 ...... .
July 29 ...... .
AUg. 5 ..... ..
Aug. 12 ...... .
Aug. 19 ..... ..
TABLE 15.
Depth Depth Depth Depth Depth Depth pepth Depth
Oto6 6to12 lto2 2to3 3to4 4to5 5. to 6 6to7
inches. inches. feet. feet. feet. feet. feet. feet.
Wilting Wilting Wilting Wilting Wilting Wilting Wilting Wilting
ecefli- coefIi- coefli- coefli- coefli- ecefli- ecefli- coefli-
cient 7.9. cient 8.4. cient 4.9. cient 5.1. cient 6.5. cient 6.8. cient 8.7. cient 7.0.
0.6
-1.0
-0.5
-3.5
-2.8
5.0
-2.0
-4.7
-4.0
-3.4
-0.5
-0.5
-1.0
-3.3
-2.4
0.8
0.0
-3.2
-4.0
-3.0
1.8
3.';
3.2
1.2
-0.6
1.1
1.8
0.4
-0.4
0.3 -0.8 ......................
3.9 2.0 0.7 -0.4 0.3
2.5 1.1 1.6 1.9 4.4
-1.0 -0.8 -0.6 0.5 1.8
long series of soil-moisture determinations, coupled with a dynamic
study of root growth through at least one wet and one dry cycle, can
answer this question of moisture penetration and its correlation with
root development.
In the short-grass community at Akron, Shantz found that for a .
period of several years little or no water was available for plant growth
below a depth of 18 to 24 inches. Even a rainfall of 2.4 inches in a day
had no effect upon the soil moisture below 18 inches. He states that
H almost the entire root system of short grasses is limited to the sur-
face 18 inches," while H deep-rooted plants are not found here for the
reason that at greater depths the soil contains no water available for
their use" (1911: 38). The results obtained by the writer in an area
of short-grass land locally known as "adobe" do not correspond with
these findings. Careful examination was made of the root systems of
Bouteloua gracilis and M uhlenbergia gracillima in pure short-grass land
about 25 miles southeast of Colorado Springs (plate 20, A). Here
Bouteloua gracilis was dominant with Opuntia polyacantha, while M uhlen-
bergia gracillima was very abundant in matlike areas. Schedonnar dUB
paniculatus, Senecio aureus oblanceolatus, Erigeron pumilus, Gutierrezia
sarothrre, and M unroa squarrosa were present, but were not at all
abundant. In fact, only 15 to 25 per cent of the soil was covered
with vegetation. Roots of Bouteloua gracilis were found to be very
abundant to a depth of 40 inches, while several roots were traced to a
depth of 51 inches. M uhlenbergia gracillima roots were very abun-
dant to 50inches, and some reached a depth of 56 inches.
These findings are so different from those of Shantz that further
work seems necessary for a thorough understanding of plains root
systems. This also seems to be true for the roots of sandhill species
(p.68). Comparisons of the root habits of these plants with those of
other communities are based upon the data made available by these
studies, and may need revision when other regions are worked. The
great variations of many root systems under different-edaphic environ-
ments is clearly shown in the section on polydemics (p. 110). However,
it is certain that plains plants grow in a soil of low and
of very hard texture. The soil in the area where these studies were
made consisted of a light-colored loam intermixed with some sand.
It is spaded with extreme difficulty and a hand-pick removes it so
slowly that in digging the trenches a large pick was kept in constant
use. Therefore, roots were unearthed only with great labor. For
example, a single specimen of Argemone platyceras occupied the atten-
tion of two persons for 1.5 days, while it required the services of a
third to do the sketching as the roots were uncovered. At depths
varying from 6 to 10 feet a layer of sand occurred. This was in
good tilth condition and usually moist enough to hold when pressed
.J firmly into a lump.
The well-developed system of wide-spreading laterals, so charac-
teristic of species of Bouteloua, Gutierrezia, Artemisia, and many others,
is undoubtedly a response to the moisture in the surface soil resulting
from summer showers. Such roots would be of small advantage to
prairie plants in the Pacific Northwest. In the prairies of Nebraska
they are probably not developed so extensively because of the greater
available water in the deeper soils. This matter is further discussed
on pages 79 and 88.
A continuous record of the soil temperature was obtained at a depth
of 4 inches among these roots from June 5 until August 5. At this
time the bulb of the soil thermograph Was lowered to 8 inches. The
most interesting fact disclosed was the extreme variation in tempera-
ture during the 24-hour period. This ranged from 60 or 70 to 90
or 9.5 F. or even more. During August, at a depth of 8 inches in the
soil, the daily fluctuation was usually about 10 F. within the range of
70 to 85. The daily fluctuation of air temperature among the plants
was usually about 35 to 40 F., the air reaching a maximum of 90
or 95 F. in the shade in the afternoon and falling to 50 or 60 F. in
the morning. Table 16 gives the mean monthly temperatures for
Colorado Springs. -
TABLE 16.
Time. Temp. Time. Temp.
Jan ... 28.00 F. Aug ... 66.9
0
F.
Feb ..... 29.8 Sept .. 59.3
Mar .. 37.0 Oct .. 48.4
Apr ..... 44.8 Nov .... 37.5
May ... M.l Dec ... 30.6
June .... 63.4
July .. 67.9 Aver ... 47.3
The humidity is low. During clear days, excepting early
in the morning, it is usually less than 50 per cent and it not infre-
quently drops to 10 or 15 per cent and sometimes lower. Even at
night, when the air i'! coldest, it seldom exceeds 80 or 90 per cent, and
dew rarely forms. There is usually steady wind movement on the
plains. Records from a standard anemometer placed just above the
plants at a height of 0.5 meter show an average daily wind velocity of
120 miles during the growing season of 1918. Likewise daily water-
losses from non-absorbing atmometers placed at a height of 10 cm.
above the soil surface are relatively high. From the data given in
table 17, which shows the average daily evaporation on the plains,
it may be seen that the plains habitat is xerophytic as regards both
air and soil environment. .
In order to visualize clearly the actual root position and the con-
ditions of competition, the accompanying bisect (plate A) was made.
Similar bisects are also given for other plant communities. The
method used is the same in all cases. A representative meter-quadrat
was selected which showed both dominant and subdominant species.
After charting and photographing, a trench was dug in front of the
quadrat, care being taken to have the wall corresponding with a side
, ,
1
j
~ I
r i
I
i
I
PLATE A
p 1
,
,
Quadrat-bisect showiug the root distribution of certain dominant and subdorninant plains species:
A, Aris/ida purpurea; B, Buutelm.ta gracili8; Ar, Artemisia lrigida; P, P8oraJ.ea tenuijlqra; C,
Chrysopsi8 viUo8a; Y, Yucca glauca.
of the quadrat cut smooth and perpendicular. Then by the use of a
hand-pick the soil was carefully removed from the roots in the trench
wall along the edge of the quadrat to a horizontal distance of 4 inches.
While this was under way the roots were carefully measured and
TABLE 17.
Time.
Evapora-
Time.
Evapora-
tion. tion.
c.c. c.c.
June 7-10 .......... 52.2 July 15-22 ....... 32.3
June 10-17 ......... 69.4 July 22-29 ......... 44.3
June 17-24 ......... 35.3 July 29 to Aug. 5 ... 64.5
June 24 to July 1 ... 66.1 Aug. 5-12 ......... 49.5
July 1-8 ........... 45.2 Aug. 12-19 ......... 41.0
July 8-15 .......... 26.5
drawn in pO<lition to scale. This gives a picture of the exact root
distribution in a block of soil a meter long, 4 inches wide, and 7 feet
deep, in this particular case. Dotted lines indicate the exit or entry
of roots from or into this soil area respectively. This plains bisect
shows the complete occupancy of the soilby roots and the intense com-
petition for moisture and soil solutes that must result.
..
68 THE :ECOLOGICAL RELATIONS OF ROOTS.
V. mE SANDHILLS SUBCLIMAX.
In order to compare root development more extensively under
different environmental conditions, work was continued in a sandhill
area of Colorado adjoining the plains community and located about
40 miles southeast of Colorado Springs. Since general climatic con-
ditions are almost identical with those described for the plains, any
differences in root development may be attributed to edaphic causes.
An excellent description of the sandhill communities, considered in the
light of their successional relation, has been given by Pool (1914).
Shantz (1911) also gives a thorough treatment of Colorado sandhill
vegetation from the standpoint of its indicator significance in regard
to crop production. The general character of the sandhills and their
characteristic vegetation is shown in plates 20, B, and 21, A.
Redfieldia 1:lexuosa.-This grass is at home in the sandhills. Indeed, it is
the most abundant and controlling species of blowout pioneers. While it
may be mixed with other pioneers, it is often the only plant present in such
situations. The usually sparse and rather small clumps are connected by
means of very long, coarse, tough rhizomes, which sometimes reach a diameter
of 4 to 5 mm. but are usually smaller; they are frequently many feet in length
~ d may be traced for a distance of 20 to 40 feet on the surface where the sand
has been blown away from them. Because of the shifting sand, the depth at
which they occur is variable. Living rhizomes with vertically descending
branches were found at a depth of 38 inches and they are rather abundant
between this depth and the surface, some running horizontally, others
obliquely and sometimes almost vertically.
From the nodes of these tough rhizomes, which are from less than an inch to
6 inches apart, whorls of roots arise in addition to sharp-pointed bude. These
consist of 2 to 10 roots, but are usually 3 to 5. The roots vary greatly in
length, diameter, and direction of growth, depending upon the age and posi-
tion of the rhizome. Those near the tip of the sharp-pointed, much elongated
rhizome may be less than 1 inch in length, very fleshy, 2 to 4 mm. in diameter,
and practically destitute of branches. Others a foot or two from the tip are
12 t ~ 20 inches long and are covered with a fine absorbing system of short
rootlets, except for the 4 to 6. inches of the growing end. The older roots,
which are well branched to the very tips with much divided laterals 1 to 3
inches long, reach a maximum depth of 56 inches. They are often 2 to 4 mm.
in diameter. They not only run rather vertically downward, but also diverge
at all angles, even to the horizontal. Long, well-branched roots were traced
to distances of over 3 feet from the base of the plant at depths of 4 to 8 inches.
The laterals from these frequently ascend vertically upward and end in well-
branched termini only 2 to 3 inches below the surface. Thus the length,
position, and abundance of this rather coarse root system, together with the
rhizomes, equip Redfieldia jleruosa in an effective manner for life in the
shifting but moist soil of the sand-dune (fig. 23).
Calamovilfa longifolia.-The sand-reed is also an efficient sand-binder.
It is frequently found associated with Redfieldia jlexuosa, Psoralea lanceolata,
and other pioneers in the shifting sands of the blowout, but normally occurs
with Andropogon hallii and others somewhat later in the succession.
It forins a veritable mat of roots and rhizomes to a depth of 34 inches. A
wide trench 6 feet long and over 5 feet deep was dug on a small dune which was
THE SANDHILLS SUB CLIMAX. 69
well ca.ptured by a dense and nearly pure growth of this sand-binder. This
plant ha.s a great abundance of tough, wiry, and very much-branched rhizomes,
2 to 4 mID. in diameter, which form an underground network connecting the
apparently isolated plants. These rhizomes are thickly covered with long
Bcales and tipped with buds about an inch in length with very sharp, hard
points. This branched network of rhizomes may be formed in the soil at al

:s
t

depths from the surface to 3 feet, the deeper ones undoubtedly haVing been
buried by wind-blown sand. From these rhizomes originate multitudes of
tough, wiry roots, 1 to 2 mm. or less in diameter, which penetrate the soil in all
directions from vertically downward to horizontally. The8e roots taper
gradually and many of them reach a maximum depth of 55 to 60 inches.
From a depth of an inch to their very tips they are abundantly supplied
with laterals which extend out more or less at right angles to a distance of 1 or
2 inches or even more. These laterals are well provided with finer branches
to the third and fourth order. Since these roots arise not only from the surface
but also from the deeper rhizomes and penetrate to 3 or 4 feet in all directions,
they form a confused tangle. Where the grasses are at all thick, as shown in
plate 21, B, the roots completely fill the soil and bind it so thoroughly that a
vertical face of a trench 6 feet long and nearly as deep held tenaciously with-
out caving, even after it was somewhat undercut at the base.
Andropogon hallii.-This tall, coarse grass forms loose, open bunches with
only a few large stems. It is a dominant along with A. scoparius in the bunch-
grass subclimax of the sandhill region. Successionally it occurs somewhat
later than the preceding sandhill grasses. It was excavated on a partially
captured sand-dune (plate 22, A). It is a very coarse, glaucous grass con-
nected by an elaborate system of rhizomes 3 to 4 mm. in diameter and lying
at a depth of 2 to 6 inches. From the rhizomes many roots take their origin,
some of which spread laterally, others penetrate almost vertically downward,
while still others take an oblique course downward. The diameter of these
varies from 1 to 2 mm. The roots running off laterally take a course almost
parallel with the soil surface or in some instances curve downward and then
up again to a point near the surface. These laterals lie at a depth of 2 to 16
inches and the maximum spread found was 3 feet. Of the roots taking a.
vertical course the maximum depth found was 27 inches; most of them,
however, penetrated to a distance of about 22 inches or less. The oblique
roots were of about the same length, but since they pass off at an angle the
depth depends upon the degree of divergence from the vertical. All roots
throughout their course were copiously branched with mostly short branches
ranging from 0.5 to 3 inches in length; as many as 8 of these branches were
counted on a single inch of the root. These small branches were themselves
abundantly supplied with minute rootlets 1 to 5 mm. in length. Occasionally
the main roots gave off a. branch 4 to 8 inches long, but these were very rare
and when they did occur were themselves branched in the same manner as
the main roots. Altogether, this plant has an excellent soil-binding and
absorbing system throughout the soil area of each bunch. Eight or nine
plants were examined.
Muhlenbergia. pungens.-This grass is characterized by tufted stems and
glaucous, narrow, rigid leaves. The culms, which arise from rootstocks, are
grouped in small tufts or cushions that lie close to the sand. It is a regular
component of blowout communities and is not infrequently dominant.
This important sandhill grass was examined on the rim and grassy top of a
blowout where it was growing very abundantly. Clusters of roots arise from
the short rootstocks, which are 2 to 6 inches long and 1 to 3 mm. in diameter.
These vary in number according to the length of the rootstock, from only 4
or 5 to 25 or more (plate 22, B); the largest are only 1 mm. in diameter. While
some of the wirelike tough roots penetrate rather vertically downward to a
maximum depth of 33 inches, others run off obliquely at various angles, even
to almost parallel with the soil surface. The lateral spread varies from 10
to 25 inches on either side of the plant. Because of the dryness of the surface
.. ch of sand, the glistening white roots are rather destitute of .branches, but
:low this level all of the roots are densely covered with multItudes of very
fine absorbing laterals. Indeed, a inch of average root w;as found to
have approximately 75 of these harrlike absorbmg laterals. While many of
them are only a few millimeters long, others reach a length of 6 to 10 inches.
All are profusely and minutely branched, the larger ones to the third and fourth
order Thus this wonderfully efficient root system is able to penetrate all
of the sandy substratum and to extract the available water., while
at the same time it forms a very effective means of preventing sand from
blowing.
Sporobolus eryptandrus.-:-In aerial habit this grass is not unlike Caia-
molJilfa. It ranks as a dominant among sandhill grasses. Three clumps of it
were examined in two separate trenches on a half-captured blowout. As a
whole it is characterized by an extremely fine fibrous root system, the main
of which arise from a short stocky rhizome 2 to 4 inches in length.
On a single rhizome 1.5 inches long, 40 of these main roots were counted. As
they leave the rhizome they pass off in all directions, some vertically, some
obliquely, and a great many of them almost parallel with the surface, but
gradually growing deeper as they pursue their course, so that at a maxi-
mum distance of 18 to 20 inches many were found at 6 to 8 inches below the
surface. The maximum vertical depth of penetration found was 22 inches.
From the point where the major divisions of the roots leave the rhizomes,
they are covered with a dense growth of root-hairs which hold the sand finnly
and give the roots the appearance of being much greater in diameter than they
really are. They are actually threadlike, usually less than 0.5 mm. in diameter
and very tough. These roots are slightly, if at all, branched for a distance of
2 or 3 inches from the rhizome, but from that point on to the very extremity
they are well supplied with branches from an inch or less to 8 or 10 inches long.
These branches pass out in all directions from the main root and are them-
selves exceedingly finely branched and rebranched to the third and fourth
order, so that the soil for a depth of from 10 to 13 inches is well filled with this
fine, delicate absorbing system.
Eriogonum microthecum.-As shown in plate 21, A, this Eriogonum fre-
quently controls local areas on sandy slopes and occurs rather widely through-
out the sandy plains area. The plant has a multicipital stem, which at a
depth of 4 to 8 inches merges into a strong tap-root 0.5 to 1 inch in diameter.
At a depth of 2 or 3 inches the tap-root begins to branch, and for a depth of
2.5 to 3 feet sends off an extraordinary absorbing system (fig. 24). Some of
these branches arise singly, but it is common to find 2 to 4 springing from
approximately the same point. In diameter they vary from 1 to 5 mm. The
general tendency of the branches is to pursue an almost horizontal course from
a distance of a few inches to 2 feet and then turn almost vertically downward,
penetrating the sand to a depth of 18 inches to 3 feet. A few of these branches,
however, instead of taking this course, pursued an almost horizontal course to
their very tips, which were sometimes 3 feet from the tap. These laterals for
the first few inches of their course as they left the tap-root were very sparsely
the distance depending upon the size of the root. Beyond that
pomt the branches gradually became more numerous and as one approached
the. extremities the number was so great as almost to form a conspicuous
network in the light sand. As the main laterals and their larger branches
approached their lower extremities, the little rootlets were so numerous as to
form fairly brushy tips. Below 3 feet the branches of the tap were much
fewer in number and much larger in size. As a rule they seem to occur in
72
THE ECOLOGICAL RELATIONS OF BOOTS.
FIG. 24.-Eriogonum microthecum.
FrG. 25.-Root system of a ten-year-old Artemisia filiJolia.
groupS, ranging from 2 to 5 in diameter. The larger after
diverging somewhat from the mam tap, took a downward course snnilar to that
of the main root. At 6 feet the tap divided up into a group of 3 or 4 roots
of almost the same size, which after diverging took the usual downward course
and at a depth of 10 feet were 2 to 3 mm. in diameter. At this depth repeated
caving of the sand made further digging so dangerous that the work was
abandoned.
Artemisia. filifolia.-In the sandhill mixed association, aside from the two
grass dominants, CaZam.oui:lfa and AndrO'J!ogon hallii, the is the most
conspicuous plant. This 18 due both to Its shrubby habIt and Its great abun-
dance. It is indicative of a light type of soil with considerable moisture pene-
tration. It drops out in the more compact hard lands of the plains. '
Three plants were examined. They were growing on a stabilized area near
the sand-dune. The largest had a tap-root 1.5 inches in diameter, from which
arose a large number of stems to a height of 2 feet, forming a bush about 18
inches in diameter. The strong, vertically descending, woody tap-root
tapered gradually and uniformly to a depth of 32 inches, at which point it
broke up into 3 laterals, the largest 0.5 inch in diameter and the other two but
slightly smaller. Some of these were traced to a depth of 9 feet, where they
were sti114 mm. in diameter (fig. 25). In the surface 30 inches of soil the tap
gave off 6 large laterals, from 6 to 13 mm. in diameter, and a very large
number of smaller ones, some of them but 0.5 mm. thick. These branches
formed a rather distinct sw'face absorbing system, and showed a strong
tendency to run out in a direction parallel with the surface. In fact, some
of them terminated in soil which was shallower than the depth at which they
originated. These with their branches and the numerous rootlets from the
somewhat oblique larger roots formed a dense network confined to the first
30 inches of soil. This was characteristic of all the plants examined and was
not unlike the root habit 'of Eriogonum. Some of these larger branches ran
off laterally to distances of 2 to 5 feet before turning downward. Although
they branched somewhat freely at intervals in the fourth to seventh foot of
soil, they often pursued their tortuous courses for long distances, sending off
hardly any laterals.
The relative abundance of the shallower roots as contrasted with the deeper
ones, together with their lateral spread and the extent of their branching, is
well shown in figure 25. Unfortunately, because of the loose texture of the
soil, it was unsafe to trace them to greater depths. The roots are dark brown
in color, the older portions having a rough, fissured bark. The younger ones,
and especially those in the deeper soil, were exceedingly brittle. The plant
here described was 10 years old. The lower branches and divisions of the tap
were very sparsely branched, the few rootlets being of the same character as
those nearer the surface. From the description and figure it will be seen that
while this species has a strong tap-root, it also has a highly developed absorb-
ing and binding system characteristic of the general group of sandhill plants.
Tradescantia virginia.na.-This spiderwort is rather common in the sand-
hills. The plants were examined on a partially captured blowout. They had
a rather fleshy root system. As many as 18 of these fleshy roots, varying from
1 to 2 mm. in diameter, were found arising from the base of a single stem (fig.
26); some passed down almost vertically to a maximum depth of 19 inches;
others started down more or less vertically, then with a gradual curve out-
ward ended from 6 to 8 inches from the vertical; others passed more or less
obliquely downward from the stem. The greater part of these fleshy roots,
however, formed a surface absorbing system which spread out in all directions
from the base of the stem, running more or less parallel with the soil surface
at a depth of 2 to 5 inches and to a maximum distiUlce of 30 inches.
All the roots, whether superficial or deeper, were practically free from
branches for a distance of 4 to 6 inches. From that point on they began to
branch, the branches increasing in number toward the tips. These branches
varied from a few millimeters to 3 or 4 cm. in length, and at intervals on these
FIG. 26.-Tradescantia virginiana.
were found short hairlike branches 2 to 5 mm. in length. While the root
system of this plant was fairly well branched, the branches were in the main
not nearly so numerous as those of most of the other sandhill perennials. The
roots were light brown to white in color, fleshy and brittle. Six plants were
examined.
Heliotropium convolvulaceum.-This annual frequently forms small fami-
lies or colonies on the rims of blowouts and on half-captured sandy slopes. The
plants were examined on August 1. They were quite mature and in full
bloom. None was over 7 inches high. They are characterized by a small
tap-root, seldom over 2 or 3 mm. in diameter, which maintains its dominance
in spite of the abundant laterals and reaches depths of 18 to 25 inches.
Throughout its rather vertically downward course, after passing through the
surface inch of dry sand, it continually gives off laterals, both short and long,
to the very tip. While many of these exceedingly fine branches do not exceed
a few centimeters in length, others with a diameter of 0.3 to 0.5 mm. extend
for a distance of 6 to 20 inches, often in a rather horizontal direction with the
soil surface. As a whole they are poorly supplied with smaller rootlets. Some-
times these run out obliquely for a distance and then turn downward. A
single root may have 4 to 6 of these long laterals. They branch at intervals
into long threadlike branches, which are themselves only moderately well
supplied with absorbing rootlets. However, the lack of abundant branches,
such as occur in many grasses and dicotyledons, is offset by the delicacy of the
root system, all parts of which are efficient absorbers. These herbaceous
roots are rather tough; they are light tan in color.
Petalostemon villosus.-This low, finely branched plant with its dense cover
of silvery leaflets stands out as a prominent object in the sandhill flora, where
it regularly plays the role of a pioneer in the blowouts. Sometimes it occupies
such situations to the complete exclusion of other species. Of the half-dozen
mature plants examined, all had strong tap-roots, about 1 cm. in diameter, a
much-branched root system, and a root penetration of 4 or 5 feet. Figure 27
is illustrative of these. An examination of this figure reveals the numerous
urface laterals with their large nodules 1 Dllll. wide and 2 to 3 Dllll. long,
~ h e i r wide lateral spread, sometimes to a distance of 2.5 feet, as well as the
bundant system of well-branched absorbing laterals. Laterals of various
:izes and often with a spread of more than a foot, come off at all depths to
near' the tip. Characteristically these run off somewhat horizontally and then
turn down rather abruptly. All of the termini are furnished with a profuse
network of finely branched rootlets. In color the roots vary from light yellow
and deep orange to brick-red. In diameter the tap was variable, lower por-
tions often being of larger size than other parts above.
FIG. 27.-Root system of PetaW8temon uillo8'U8.
Gilia longiflora.-Three plants of this annual, growing on a well-covered
sandhill, were examined, all were mature and in full bloom; their underground
system started with a strong tap-root, varying in size from 7 to 12 Dllll. in
diameter. The tap penetrates almost vertically downward, tapering very
rapidly until, at a point 6 inches below the surface, it is usually not more than
3 Dllll. in diameter. From this depth it tapers more gradually, pursuing a
kinky and tortuous course downward through the sandy soil to a maximum
depth of 40 to 50 inches (fig. 28). The most marked characteristic of these
tap-roots is the remarkable number of both large and small laterals thrown
off within the first 2 to 10 inches of soil. From 8 to 14 of these are often found,
76 THE ECOLOGICAL RELATIONS OF ROOTS. -
varying in diameter from 1 to 3 mm., not including numerous smaller ones.
These larger laterals usually pursue a course almost parallel with the surface
for a distance of 5 to 22 inches, and then almost invariably turn abruptly
downward, reaching depths of 10 to 35 inches. Throughout their course they
branch freely into both large and small rootlets, the larger branches spreading
widely and then turning downward after the fashion of the main laterals.
Throughout the course of the laterals and their larger branches, a striking
FIG. 28.-Gilia longiflora.
system of fine absorbing rootlets, branched to the second and third order,
is found. Below the point where the tap throws off the larger laterals, it is
much more poorly branched, the branches being relatively much smaller and
shorter, but well supplied with fine rootlets, as are the laterals above. From
the figure and description it will be seen that although this is an annual, the
absorptive system is characteristic of that of most of the species found in the
sandhills. The roots are glistening white and quite tough for herbaceous plants.
Euphorbia petaloidea.-Euphorbia is a sandhill pioneer which forms both
families and colonies. The root system begins with a strong, smooth tap-
root, varying in diameter according to the age and size of the plant, the
THE SANDHILLS SUBCLIMAX. 77
FIo. 29.-Underground parts of Psoralea lanceolata, showing root tubercles at a
depth of 8 feet.
78
largest examined being 4 mm. in diameter. This tap penetrates almost
vertically downward. Beginning at a depth of 2 to 3 inches, it sends off
multitudes of fine fibrous roots which are much branched and penetrate the
soil on all sides to a distance from a centimeter or less to 3 or 4 inches. In
addition to these finer roots, the tap sends off longer laterals which traverse
the soil in all directions, some taking a course almost parallel with the sur-
face, while others pass off more obliquely. These vary in length, some being
10 to 12 inches long. The laterals are much branched andrebranched, and
together with the finer rootlets form an effective absorbing system. Usually
at a depth of 5 to 8 inches the tap divides up into such a system of large
laterals as to almost if not quite lose its identity. These lower laterals vary
from 0.5 to 1 mm. in diameter and diverge more or less obliquely downward,
some penetrating to a depth of 26 inches or more. The deeper penetrating
divisions of the tap branch rather freely and end in much divided termini.
On the whole, however, these lower branches are more sparsely supplied with
absorbing rootlets than those nearer the surface. The roots are of a .light
brown to white color, are rather fragile, and hard to trace. Six plants of
varying sizes were examined.
Psoralea lanceolata.-This legume sometimes replaces Redfieldia ftexuosa
as the pioneer in the shifting soils of blowouts, but normally it belongs later
in the succession. Its underground parts eminently fit it to succeed in a
habitat where the ground-line is constantly lowered by wind erosion or built
up by the same agency.
The individual plants, which may be a few inches or several feet apart, are
connected by a more or less horizontal system of rhizomes, which varies in
depth from 2 or 3 inches to more than 2 feet. These connecting parts vary
from a few millimeters to more than a centimeter in diameter and may run
horizontally for distances of 10 to 30 feet or more. At irregular intervals
along their course occur erect portions, often a centimeter in diameter and
frequently forked at the top. These give rise to the individual plants, while at
other intervals, and not always below the erect stems, strong, rather vertically
descending roots occur, many of which reach a depth of 8 or 9 feet (fig. 29).
The surface 2 or 3 feet of the root system is poorly supplied with absorbing
rootlets. In fact, only a few of the major branches have their origin' in this
layer of soil and the ultimate rootlets of these extend much deeper. The main
system of branches begins in the fourth foot of soil and extends to the 8 or
9-foot level. This consists of both large and small wide-spreading and well-
branched rootlets, the laterals often running out to a distance of 2 feet or more
from the base of the tap. Large root nodules, 1 to 3 mm. in diameter, were
observed at a depth of 8 feet. One large tap-root was traced to a depth of
over 9 feet, where it still maintained a diameter of 4 rom. It was not followed
further, because of the danger of caving the sandy-walled trench. The roots
were dark brown in color and rather tough.
Ipomooa leptophylla.-The bush morning":glory is a common plant in the
sandhills, where the large hemispherical tops, conspicuous because of their pro-
fuse purple blossoms, cover many square feet. Two plants which were growing
together were examined. The top measured 7 feet in diameter and was 30
inches high. The multicipital stem arose from strong tap-roots 2.5 inches in
diameter. At a depth of 9 inches the taps became greatly enlarged, reaching
a diameter of 6 to 8 inches, which they maintained to a distance of about a
foot, when they tapered off gradually, so that at a depth of 4 feet they were
but an inch or two in diameter. Below this point the tap lost its dominance,
breaking up into large numbers of deeply penetrating and widely spreading
branches, as shown in figure 30.
THE SANDHILLS SUBCLIMAX. 79
In the first foot of soil scarcely any laterals originated, but below this point
both large and small branches came off in great abundance. These varied in
diameter from less than 1 mm. to half an inch. While a very few ran some-
what vertically or obliquely downward, the general course of most of them
was quite parallel with the soil surface. The lateral spread was enormous,
the roots running off to distances of 15 to 25 feet or more. One of the larger
forks of a root, 0.5 inch in diameter, was traced to a distance of 14 feet and at
a depth varying from 25 to 30 inches from the base of the plant, when it
branched into two equal parts, 4 mm. in diameter. These wide-spreading
laterals sent off abundant branches, many of which passed off at right angles,
either vertically downward or vertically upward, for distances of many feet.
Indeed. the surface foot of soil as well as the 10 feet below it were literally
filled with the glistening white, brittle branches of this remarkable root system ..
The ultimate branches, however, while occurring in great numbers, were not
profusely divided. The great network of roots originating at a depth of 5 feet
is well shown in figure 30, but because of the caving of the sandy soil it was
impossible to follow them to a greater depth. However, judging from their
diameters of 2 to 4 mm. at this level and the nature of the other roots examined,
it is highly probable that they penetrated many feet deeper. The enlarged
portion of the tap-root not only furnishes an enormous reservoir for food, but
also a storehouse of water upon which the plant may draw during a period of
drought.
SANDHILL ROOT SYSTEMS AND 1HE SANDHILL ENVIRONMENT.
Eight of the 19 sandhill species examined
l
have roots which are
entirely or nearly confined to the first 2 feet of soil. Of the others, all
but one have their main root development in the first 2 or 3 feet of
soil. Even the very deep-rooted Eriogonum microthecum and Arte-
misia jiZifolia show a striking profusion of long, wide-spreading laterals
in this soil layer. Indeed,it will be shown later that Stipa comata,
BouteZ.()ua gracilis, and Chrysopsis viZlosa, all rather deep-rooted plains
species, become rather shallow-rooted under sandhill conditions.
Among the grasses the rhizome habit is extremely well-developed,
while long, rather horizontal, shallow laterals are characteristic of
most of the plants examined. In this soft substratum such a root
habit can not be attributed to the mechanical difficulty of penetration,
but seems closely related to the water-supply.
Rainfall is at once absorbed and there is practically no run-off, even
in the heaviest showers. As soon as the storm passes, evaporation
dries out the surface sand with great rapidity, but to a slight depth
only. This surface layer of dry sand forms an excellent mulch, which
has a wonderfully retarding effect upon further evaporation. At a
depth of but a few inches below the surface the sand is always moist
and may usually be molded into lumps by pressure of the hand. The
vegetative cover is usually rather sparse and consequently water-
losses by transpiration should be much less than on the plains. U nfor-
tunately few data on soil-moisture in sandhill soils are available.
1 Only 14 are here described; the others will be found under the section on ecads (p. 110).
80
Bates (1910) gives us the following table of water-contents for soils
taken in the sandhills at Halsey, Nebraska:
TABLE 17.-Amount of moisture in the soil, per cent of dry weight. ~ 1 I ! i I r ~
At 1 foot. At 2 feet. At 3 feet. At 6 feet.
Station.
May. July. Sept. May. July. Sept. May. July. Sept. May. July. Sept.
1---------------------
South slope ..... 4.1 1.9 3.1 4.8 1.6 3.1 4.3 1.8 3.2 3.9 2.8 3.3
Bottom ......... 5.1 2.9 4.7 4.8 2.1 4.2 5.4 3.3 4.2 6.5 7.4 8.6
North slope .... 0.1 3.0 4.7 5.3 3.3 5.0 5.9 0.1 6.7 0.9 7.3 7.4
Ridge .......... 3.3 4.0 3.6 4.4 3.8 3.9 5.2 3.7 3.7 0.4 4.2 4.9
From these data it may be seen that the soil was rather uniformly
moist to a depth of 6 feet. We must keep in mind, however, that these
data were taken under a rainfall of about 23 inches and during the wet
phase of the climatic cycle.
A single set of duplicate determinations, taken in the area studied
on July 30, showed the water-content to be rather uniform to a depth
of 3 feet (about, 4 per cent), 3 per cent of which was available for plant
growth. While further determinations were not made, in all of the
numerous trenches dug the sand was perceptibly drier at greater
depths. It seems certain that under the normal light rainfall the
surface soils from 4 to 36 inches would contain the most moisture,
while the deeper soils would become wet only during the years of
abnormal precipitation. The latter may account for the deep root
habit of Psoralea lanceolata and others. The deeper soils, once wetted,
would dry out very slowly, because of the relatively small number of
plants drawing their water-supply from them. Soil nutrients may be
a limiting factor, but this seems improbable in view of the luxuriant
growth and complete occupancy of the soil by plants when sufficient
water is supplied.
THE GRAVEL-SLIDE COMMUNITY. 81
VI. THE GRAVEL-SLIDE COMMUNITY.
A fine series of diverse habitats for investigation are found in the
mountains adjoining the plains. During August a detailed investiga-
tion was made of the roots of plants at an altitude of about 8,000 feet
in the Pike's Peak region of the Rocky Mountains. Here the soil is
composed of disintegrated granite, the degree of disintegration and
decomposition determining largely the type of plant community
occupying any particular area. The successional sequence and species
belonging to each community of the sere may be found in publications
by Clements (1904, 1905). The first herbaceous plants to occupy the
new soils formed by the crumbling granite on the steep mountain slopes
are members of the gravel-slide community (plate 23). The semibare .
areas thus populated are so extensive and the life conditions so unique
that a rather detailed study was made of the most important species.
Krynitzkia virgata.-This striking plant has an erect, spike-like stem seldom
reaching a height greater than 8 to 12 inches in this habitat. It is a principal
species of the gravel-slide community. It has a tap-root 4 to 9 mm. in diam-
FIG. 31.-Krynitz1cia virgata. The heavy slanting line indicates the actual ground-line.
eter, which reaches depths varying from 20 to 30 inches (fig. 31). Except for
the first 3 to 5 inches, where the plant tops have slipped down the gravel
slide, the root pursues a rather vertically downward course, the tip being only a
few inches horizontally away from the base of the plant, in spite of the back-
ward and forward meanderings of the root through bends of 3 or 4 inches in
diameter. The tap is often flattened and kinked where it forces its way
through the crevices of the semi-decomposed rock. The number of larger
branches is few, usually not exceeding 2 or 3. One plant gave off only one
large branch, about 2 mm. in diameter, at a depth of 3 inches. AB is charac-
teristic of numerous other plants examined, this lateral branched freely and
ran off in a direction nearly parallel with the surface for about 15 to 20
inches, ending in a network of well-branched rootlets. On another plant a
lateral ran off more than 36 inches from the base of the stem, branching again
and again. Other smaller branches arise from the tap in great abundance.
These are only a few centimeters long, but well-branched and densely covered
with root hairs. The older cortex is black and of a papery texture, peeling off
readily. .
Paronychia. jamesii.-This small, cespitose perennial forms a consocies of
the community. It has a tap-root about 5 mm. in diameter. Like all other
plants on the gravel-slide, the top has been pushed down the slope several
FIG. 32.-Paronychia jameBii.
inches. The tap breaks up into several larger laterals, usually 3 to 5, and a
great number of smaller ones. . Some of the laterals run off in a horizontal
direction to a distance of 30 inches or more, at about 2 to 4 inches in depth,
branching repeatedly, as shown in figures 32 and 33, and being abundantly
u lied with minute, absorbing rootlets. Others run off obliquely, while
StiliPothers parallel more or less the attenuated tap-root, and like the tap are
:bundantly supplied with both long and short rootlets, the whole forming a
wonderfully efficient absorbing system. Compared with the transpiring sur-
face the root system is very well developed. The surface view in the figure
the roots at a depth of 2 to 2.5 inches. The dotted lines show where
they turn downward. No roots were found to penetrate to a greater depth
than 30 inches.
Aletes acaulis.-This low umbellif-
erous plant, which is only 6 to 8 inches
high in fruit, is the dominant of the
gravel-slide, often being more abundant
and conspicuous than all other plants
combined (plate 23, A, B). From the
large tap-root arises a large number of
stems, each multicipital in character;
47 individual stems were counted on
a single medium-sized plant. Such a
group forms a formidable obstacle to
the rock fragments moved by gravity,
the latter pushing the top of the plant
6 to 10 inches downward from the top
of the well-anchored tap-root. Both
the main stems and the branches show
marked wrinkles from profound con-
tractions. At the base of the shoots of
this perennial, great clusters of dead
and decaying gray leaf-bases remain
attached to the plant. ,
The tap-root is often 4 cm. or more
in diameter. It is dark brown in color
and quite spongy in texture, as was
noted for several species in this habitat,
the fleshy roots probably serving for
water storage. One plant with a tap 4 FIG. of a single root of
cm. in diameter, from which originated !'aronychw;amesu at a depth of 2 to 2.5
7 d
t' t te I . . h mches.
18 mc s m-c usters, gave nse ill t e
first 3 inches of soil to numerous laterals ranging from 1 cm. to only a few
millimeters in diameter (plate 24, A, B). One of these laterals, 3 mm. in
diameter, ran off at a depth of about 4 inches and in a direction parallel with
the soil surface to a distance of 4 feet, giving off numerous branches, both
large and small, each of which, after branching profusely, ended in a network
of tiny, much-branched laterals. Another surface lateral,8 mm. in diameter,
ran up the slope at an average depth of 5 inches to a distance of 4.5 feet. At
a depth of 6 inches the tap broke up into 3 parts-7, 8, and 10 mm. in diameter
respectively. These were very much curved and twisted. They followed
the crevices of the rocks and none reached a depth greater than 30 inches,
but spread laterally to a distance of 3 or 4 feet or more from the base of the
All of the laterals branched profusely and terminated in the moist rock
creVIces in networks of tiny rootlets. The surface 2 to 5 inches of soil is
especially filled with these absorbing laterals, but they are abundant through-
out the soil to the tips of the deepest roots. Plate 24, B, shows a fragment of
these branches. In fact, the soil is literally filled with these absorbing rootlets
to a point several feet on each side of the plant. A single plant thus lays hold
of the surface soil to a depth of 18 inches or more within a radius of 4 or 5
feet from its base.
Apocynum androsremifolium.-Large areas, especially of the steeper gravel
. slides, are frequently covered by extensive communities in which this species
is dominant, often forming families (fig. 34). The vertical portions ~
from the horizontal rootstock are 2 to 5 mm. in diameter. From theselorigi-
FIG. M.-Apocynum androsmnifolium, showing rhizomes and dense network of roots.
nate groups of 2 to 5 erect stems. The rootstocks lie at a depth varying from
6 inches to 2 feet, but usually at about 8 to 16 inches. These run for distances
of 6 to 10 feet or more, giving rise at irregular intervals to erect portions
which bear new plants. Sometimes these connected plants are only a few
inches apart, while at other times the interval between them may be 3 or 4 feet.
The vertically ascending parts seldom have large branches, but are well
clothed with abundant laterals 1 mm. or less in diameter. Theselbranches
divide and subdivide into many branch orders, forming brushlike mats which
run off in all directions. to a of 6 or 8 or more. Howe'!er, the
most profound branching anses from the honzontal rootstocks. Like the
former these are not large in diameter, but are branched so profusely, extend-
ing verlicallY upward as well as downward and laterally, that they com-
pletely occupy the soil from a of 2 inches to a maximum depth <?f 4
Many of these deeper roots ongmate from strong laterals 1 to 2 mm. m diam-
eter which branch off and run in various directions from the rootstocks.
34 illustrates the profound absorbing surface characteristic of these
gravel-slide plants. In color, the larger roots are dark brown, while the finer
ones vary from tan to almost white.
Smilacin80 stella.ta..-Faroilies of this plant often occur on steep gravel-
slides or they are the dominants of a gravel-slide colony. Stout stems, from
3 to .; mro. in diameter, arise at intervals of an inch to more than a foot from
the stout horizontal rhizomes. The latter are about the same diameter as the
stem and lie usually at a depth of about 4 to 6 inches, although they are some-
times shallower where the gravel has rolled away and often much deeper
where the gravel has covered them (plate 24, c). Some were found at a depth
of 30 inches. The rhizomes are much branched,frequently at right angles, a
single rhizome system often connecting a whole family of plants through a
distance of 7 or 8 feet or more. These rhizomes furnish not only an excellent
means for propagation, but also serve as storage organs for these herbaceous
perennials.
The plants frequently grow in such dense clumps that the soil at a depth of
4 to 6 inches is quite filled with dense masses of these rhizomes. Although the
vertical stems do not give rise to rootlets, the rhizomes are uniformly covered
on all sides with rootlets about 0.5 to 1.0 mm. in diameter. These run off in all
directions, including the vertical to a distance of 3 to 6 inches. Although
they are entirely unbranched, they are completely and uniformly covered with
a dense coat of root-hairs. Thus the plant is well provided for absorbing the
moisture in the shallower soil. In addition to these shorter rootlets, groups
of 2 to 5 larger branch roots arise at frequent intervals at the base of the
vertical stems or where these stems have been, a place now marked by a seal-
like scar. These vary from 2 to 4 mm. in diameter and throughout their
course are densely covered with root-hairs. They penetrate the soil to a
maximum depth of 44 inches. Through the first 6 to 12 inches of their course
they are unbranched or at least poorly branched, but from this point they
branch profusely to the third or fourth order, the branches spreading widely
and the ultimate rootlets being rather coarse, often 0.2 to 0.5 mm. in diameter. c'
Since these branches run obliquely and even horizontally as well as vertically
downward, they furnish with the shorter roots already described an excellent
absorbing system. Altogether the finer rootlets are very coarse and poorly
branched when compared with Aletes, Thlaspi, and certain other gravel-slide
plants. The entire underground parts vary from tan to white in color.
Pachylophus cmspitosus.-This is a common plant on bare gravel-slides
where it forms pioneer consocies. The social habit is due to the method of
vegetative propagation (fig. 35). The plants are frequently connected by
rhizomes from which arise roots 2 to 5 mm. in diameter, none of which were
found to reach a depth greater than 45 inches. Frequently at 4 to 8 inches
deep, they turn off abruptly up or down the slope and run parallel with the
surface for long distances. Large branches arise from these roots and do much
?f the absorbing for the plant. Of numerous specimens examined, the one
illustrated in figure 35 is typical.
Both large and small rootlets may arise from the rhizomes. The main
laterals are well supplied with rootlets and the soil is quite filled with them
to a depth of 6 to 36 inches. As compared with Aletes and Apocynum, the
rootlets are coarse and rather poorly branched. They vary from tan to nearly
white in color. The tap, as in many gravel-slide plants, is rather fleshy.
FIG. 35.-Pachylophm COlspitosm.
Thlaspi alpestre.-This penny-cress is abundant but rather inconspicuous,
being only 5 to 8 cm. high. Its tap-root is usually not over 2 Mm. in diameter,
and penetrates the dry surface gravel to a depth of 1.5 to 2 inches before it
gives off branches. Beginning at this depth, tiny laterals, ranging from almost
microscopic in size to 1 Mm. in diameter, develop in abundance (plate 24, D).
The larger laterals are relatively few, usually not more than 2 to 5, but the
smaller ones, many about the size of a fine silk thread or smaller, occur in
great abundance. Often they run parallel with the surface throughout the soil,
branching repeatedly in such a manner as to form a delicate mass of rootlets,
the ultimate termini being minute. The lateral extent is usually not greater
than 4 to 7 inches and the depth of the plant seldom exceeds 15 inches. To
really appreciate the delicacy of the profusely branched fragile root system,
it is necessary for one to carefully pick away the coarse rock particles.
Mentzelia multiflora.-Size, duration, and abundance combine to make
this species a dominant on many gravel-slides. The plants are usually about
10 to 14 inches high when in full bloom. The roots start with a tap 12 mm. or
less in diameter, which, after penetrating to a depth of 2 to 5 inches, sends off
strong laterals as large as 3 or 4 mm. in diameter. These are distinctly shallow,
run almost parallel with the surface, and are repeatedly branched, the
fine rootlets ending in hairlike extremities. Although they seldom run more
than 18 inches from the base of the tap, they are frequently much longer
because of their curved and twisted course. The tap-root takes an almost
vertically downward course, except where it follows for a time the crevices in
the rocky soil, and penetrates to a maximum depth of about 22 inches.
Although the tap is not so profusely branched as are the shallower laterals, the
branches of both are very similar. The smaller branches of both the laterals
and the tap are profusely covered with a fine growth of root-hairs. Although
this plant, when compared with most others, has an exceedingly well-developed
absorbing system, it does not compare favorably with the wonderfully exten-
sive and excellently developed root system of Aletes. However, both are well
adapted to get the water falling during the frequent mountain showers and
thrive under conditions where most plants coUld not grow. The roots are
almost white in color and very spongy in texture.
Eriogonum llavum.-This plant is a subdominant in the gravel-slide com-
munity, its masses of yellow flowers making it very conspicuous in the
autumnal aspect. The specimen had a strong woody tap-root 2 cm. in
diameter which tapered within a length of 6 inches to only 3 mm. in diameter.
Here it gave off 2 laterals each 2 mm. in width. These ran off laterally for
about 2 feet at a depth of 3 to 7 inches. An enormous number of smaller
profusely branched laterals arose, forming a dense absorbing network about
FIG. 36.-Eriogonum ftavum.
the plant for a radius of more than 18 inches. These were extraordinarily
well-branched and rebranched and were very dense. At 5 inches in depth
another large lateral occurred, and the root broke into 2 nearly equal parts at
6 inches in depth. One of these ran off into the bank and slightly upward to
a distance of 2 feet from the base of the crown, but it was really much longer.
It ended 3 inches below the soil surface. This lateral, scarcely more than 1
mIn. in average diameter, gave off both short and long branches in a profuse
manner, all being repeatedly branched into minute termini and densely covered
with root-hairs. They also dipped upward to within 2 inches of the surface
and downward to 8 or more inches, but were most abundant at 4 or 5 inches in
depth. The other root at the fork ran out into the slope for over 5 feet at an
average depth of 16 inches. The roots often run long distances without much
branching, but the last 2 or 3.5 feet form a great network of branches. The
roots are reddish in color and relatively tough. None of those examined were
deeper than 2 feet (fig. 36).
In another specimen the root system began to divide almost from the
crown, some of the main parts, however, being much larger than the others,
varying from 2 to 13 mm. All the rootlets, whether large or small, were
marked by a diffuse dichotomous branching. Most of the branches were
superficial, extending to a depth of 1 to 12 inches. These branches were
themselves branched many times, and the termini of the branches of all sorts
were persistently marked by capillary roots, ramifying and forming a con-
spicuous network in the soil. The course of most of these laterals was par-
allel with the surface. The maximum spread of the main absorbing branches
was found to be 4.5 feet. A very few of the larger branches take a more
marked downward course, branching very much the same as the horizontal
laterals, but the branches are not so numerous nor so long. The maximum
depth recorded was 39 inches and the tip of the root had wandered 43 inches
from the vertical.
GRAVEL-SLIDE ROOT SYSTEMS AND GRAVEL-SLIDE ENVIRONMENT.
All of the plants of this community are similar in possessing roots
adapted to secure moisture and nutrients from the surface soil. They
are characterized by a shallow, widely spreading root system confined
largely to the :first 18 inches of the soil and in fact usually best developed
in the surface foot. The lack of depth is compensated by a remarkably
wide lateral extent combined with a profound system of branching.
An explanation of these adaptations is readily found in a study of
the soil and the distribution of the rainfall.
The gravel-slide soils consist of a superficial layer of coarse angular
rock fragments varying in size from over an inch to a few IIiillimeters
in diameter. Except during rains this surface layer is very dry. The
slope is so steep that there is often a constant moyement of the rock
particles down the mountain-side, the course of these moving pebbles
. being marked by depressions looking not unlike the tracks of harrow
teeth. Most of the plant tops have slipped down the slope from 2 to
8 or more inches. This surface layer is very efficient in preventing
run-off, as well as in forming a dry mulch and thus protecting the
underlying soil from high evaporation. A concrete illustration of the
effectiveness of this layer is shown in the following experiment. Two
metal cylinders, 8 inches high and 5.5 inches in diameter, closed at one
end were filled with wet soil and one of them was covered with a layer
of these coarse gravel particles to a depth of an inch. Mter an equal
exposure to evaporation for a period of 30 hours, it was found that
the one container had lost 202 grams of water, which was more than 8
times as much (24 grams) as evaporated from the soil covered with
the gravel mulch. In nature, finer particles occupy the interstices be-
tween larger ones, and hence the gravel mulch must be much more
~ ~ .
Below this surface gravel are about 4 inches or more of fairly well
decomposed rock, a mixture of coarse gravel and sand. On older
TABLE 18.-Water-content of tM 80il of the gravel-8lide and half-gravel-8lide during 1918.
Depth 0 to 6 inches. Depth 6 to 12 inches. Depth 12 to IS inches.
Date.
Gravel- Half-gravel- Gravel- Half-gravel- Gravel- Half-gravel-
slide. slide. slide. slide. slide. slide.
June
8 ........ 4.7 3.0 5.1 3.3 4.2 2.8
June
11 ........... ......... 3.8 . ........ 2.7 . ........ 2.9
June
24 ............ 5.2 9.2 4.3 4.4 5.0 3.2
July 1. ........... 3.1 5.5 4.0 4.7 8.5 4.2
July S ............ 3.3 3.5 4.2 2.4 3.2 2.2
July 15 .......... 4.0 9.5 4.5 7.5 3.5 4.0
July 22 ........... 6.0 10.1 3.9 5.7 4.1 4.3
July 29 .... 4.2 6.5 4.1 3.9 3.2 3.2
Aug. 5 ............ 4.9 S.1 3.2 2.1 2.7 2.1
Aug. 12 ............ 4.5 8.9 3.0 4.4 2.6 4.3
Aug. 19 ............ 5.1 6.6 4.3 3.S 4.6 3.4
slides this layer extends much deeper. Below this the soil changes
from a dark brown to a more reddish color and consists of fairly well
decomposed granite, which becomes less broken up as one goes deeper.
At 2 to 4 feet in depth it changes into almost solid rock. The roots
show a marked tendency to follow the cleavage planes of the rock.
Below 4 to 6 inches the soil is remarkably compact, and it is necessary
to remove it with a pick, this sometimes being accomplished with con-
siderable difficulty.
An examination of the weekly soil-moisture determinations in table 18
shows that while the water-content is at no time high,it is rather uni-
formly distributed throughout the first 18 inches of soil. Owing to the
extreme irregularity in degree of fragmentation of the rock particles
and to the heterogeneous nature of these soils, moisture-equivalent
determinations are not given. The amount of non-available water
in the gravel-slide soils was found to vary from 1.9 to 7.8 per' cent;
in the half-gravel-slide from 2.0 to 8.6 per cent; and in the soils of the
forest floor from 2.7 to 10.5 per cent (page 109).
The rather high evaporating power of the air as measured by non-
absorbing atmometers is given in table 19.
TABLE 19.-Average daily evaporation on the gravel-slide and half-gravel-slide during 1918.
Gravel-
Half-
Gravel-
Half-
Date.
slide.
gravel- Date.
slide.
gravel-
slide. slide.
c.c. c.c. c.c. c.c.
June 6-10 ........ 26.6 20.2 July 15-22 ........ 19.8 15.5
June 10-17 ......... 51.6 40.1 July 22-29 ........ 31.5 21.4
June 17-24 ......... 25.8 22.8 July 29 to Aug. 5 ... 49.4 31.1
June 24 to July 1. .. 48.2 30.0 Aug. 5-12 ......... 22.9 17.0
July 1-8 ........ , 35.8 25.0 Aug. 12-19 ....... 24.2 16.9
July 8-15 ........ 17.6 13.2 Aug. 19-23 ........ 54.2 32.0
One factor greatly accelerating evaporation is the marked wind
movement. From June 6 to August 23 the average daily wind velocity
at a height of 0.5 m. was 103 miles. Notwithstanding the high evapora-
tion losses, nearly all of which occur during the day, the upper layer
of the soil containing the roots is kept moist by frequent rains. Seven-
A
I",
F r
:A
. ~ '
~
rr
f
.,.
~ t F '
FIG. 37.-Quadrat-bisect showing root relations of gravel-slide plants. This was made
along the front of the quadrat shown in plate 23, B. P, Paronychia jamesii; A, Aletu
acaulis; K, Krynitzkia virgata.
teen showers, varying in intensity from 0.1 to 1.85 inches, occurred
from June 21 to August 28, 1918, while the total rainfall during the
period was over 10 inches. As already pointed out, none of this water
runs off except during the heaviest rains, while the loose gravel inhibits
high evaporation losses. Moreover, the plants are so sparsely spaced
that only relatively small amounts of water are removed by transpira-
tion. Perhaps 95 per cent of the rocky soil surface is bare except for
Pai'melia saxatilis and a few other crustose lichens (plate 23). This
habitat is somewhat similar to that of the sandhills in respect to
the dry surface mulch and the supply of moisture in the upper layer.
The root habit is clearly a response to the environment. Undoubtedly
the large number of roots which have such a strong tendency to run
up the slope serve in part for anchorage. The distribution of soil
nutrients may alSo have some share in this. The bisect in figure 37
shows that root competition on the gravel-slide is far from severe. .
VII. TI-IE HALF.GRA VfL.SLIDE COMMUNITY.
The half-gravel-slide community represents a distinct successional
advance over that of the gravel-slide, both in the diversity of species
and the density of plant population, as well as in the correspondingly
more favorable habitat (plate 25, A). It is an intermediate stage
between the gravel-slide and the forest. Here the root systems of the
most important grasses, herbs, and shrubs were studied.
Elymus triticoides.-This grass is perhaps the most abundant and important
species of the half-gravel-slide community. Because of its excellent root
development and consequent ability to compete successfully with other
species, it not infrequently controls large areas, sometimes almost to the
exclusion of other plants.
Elymus was excavated at the half-gravel-slide station in coarse, rocky soil
(plate 25, A, B). The soil of the upper 18 to 22 inches was light brown to gray
in color. The roots were densely matted. The larger were 1.5 to 2 mm. in
diameter, being much branched into fine rootlets supplemented by countless
finer well-branched and matted ones. In fact, the soil to a depth of 15 or 18
inches was completely filled with a dense network of roots. The larger ones
penetrated to a maximum depth of 46 inches. The mass of roots at the surface
had a lateral spread of about 18 inches on either side of the bunch. Many of
the larger roots descended at an angle of about 30 degrees from the vertical
to a depth of 15 or 20 inches, when they turned directly downward to a depth
of 40 to 45 inches. All of these deeply penetrating roots were, like the others,
extremely well branched to the third and fourth order and ramified through-
out the crevices of the gravel. Some of these larger roots, after reaching. a
depth of 13 inches, took a course up the steep slope, following at this depth a
line approximately parallel with the surface of the soil. They were well
branched, the branches coming up to a distance of only 5 or 6 inches from the
surface of the ground. The lower part of the root system was white in color
and extremely fragile.
Solidago oreophila.-The autumn landscape is frequently given tone by the
masses of yellow flowers of this important half-gravel-slide species. This
plant consists of clusters of stems connected by short rhizomes, thus forming
a clump. From the base of the rhizomes arise great numbers of fibrous roots
about 1 mm. in diameter; as many as 50 to 75 may originate from a single
inch of the rhizome. Numerous roots run off parallel with the soil surface, or
nearly so, to a distance of 20 to 30 inches from the base of the plant, sending
off rather numerous branched and rebranched threadlike laterals from a few
centimeters to a few inches long and finally terminating in a much-branched,
brush-like ending. Many of the shallower roots also run off obliquely, so that
at a distance of a foot from the plant they may reach to a depth of 8 to 10
inches or even more. However, these are not so numerous as the superficial
ones. These shallower roots frequently turn down near their tips to a depth
of 6 to 15 inches.
The vertically descending roots send off laterals rather sparingly to a dis-
tance of 12 or 18 inches,. beyond which depth they become more and more
profusely branched, finally terminating in great clusters of hairlike, minutely
branched ends. In the more decomposed soils the deepest roots may pene-
trate to a distance of 30 to 38 inches, while in the more rocky substratum
they are much shallower. This deeper group of roots usually spreads laterally
THE HALF-GRA VEL-SLIDE COMMUNITY. 93
to only 6 or 8 inches on either of the b.ase of the plant. The roots are
Ii ht'tan in color and show consIderable tensIle strength (fig. 38).
g Rubus deliciosus.-This plant is one of the most important of the shrubs
of the half-gravel-slide. With Opulaster and certain others it forms the
transition stage to forest... . . .
Three specimens were exammed which were so th.at only one will be
described This plant arose from a tap-root 15 mm. ill diameter. It had 3
main two of which reached a height of 20 inches. At a depth of 3
inches the gave off 2 laterals, 2 and 3 mm. in diameter respectively.
The smaller of these ran off almost horizontally to a distance of 20 inches,
giving off great numbers of much-branched laterals, the termini being almost
FIG. 38.-Solidago oreophila, showing wide-spreading lateral and deep central roots.
hairlike and reaching a depth of over 20 inches (plate 25, c). The larger
lateral ran off in the opposite direction to a distance of more than 2 feet,
branching repeatedly and ending in clusters of rootlets only 4 inches below
the soil surface. On the first 5 inches of the tap several other smaller rootlets
occur:ed,. while at 6 inches depth the root divided into 2 equal parts about 6
ill diameter. One of these soon rebranched, while all ran off more or less
honzontally or obliquely, some to a distance of approximately 3 feet from the
base of the plant. The maximum depth did not exceed 36 inches. These
roo!S branched repeatedly into both large and small laterals, those running
honzontally being characterized by a multitude of shorter, minutely branched,
often vertically descending rootlets. Thus the plant is provided with an
effective absorbing system, which ramifies widely and fills the soil from a
depth of from 4 to 36 inches. The whole root system is characteristic of the .
half-gravel-slide root habit. ~ .'
Besseya. pla.ntaginea.-This plant frequently grows in clusters of 3 or more.
the individuals of which are connected by short rhizomes about 5 mm. or less
in diameter and 2 or 3 inches long. The base of the plant and these rhizomes
are densely covered with fleshy roots about 2 mm. in diameter. AB many as
30 to 40 of these roots occur on a single inch of the rhizome. Many of them
pursue a vertically downward course and end at a maximum depth of from
25 to 30 inches (fig. 39). Others run out rather parallel with the surface
of the soil and at a depth of 2 or more inches to a distance of over a foot, when
they turn abruptly downward, reaching a depth of 16 or 18 inches. Btill others
FIG. 39.-Be88eya plantaginea, showing the widely spreading root system.
fill in the angle between the horizontal and vertical roots, running off obliquely
and then turning downward. Very few branches occur on the first 4 to 6
inches of the root. But beyond this point the roots (which are now a milli-
meter or less in diameter) branch freely, sending off laterals to a distance of
5 or 6 inches, the last 12 to 18 inches being so well branched and rebranched
as to form a fine absorbing network. Thus the soil within a radius of at least
a foot from the plant and at a depth of from 20 to 30 inches (except the sur-
face 2 inches) is rather completely filled with absorbing rootlets of this fleshy
perennial. As its root system shows, Besseya is an excellent illustration of a
transitional forin between the gravel-slide type and that of ordinary soil.
The roots are dark tan in color and are fairly tough and resistant. Five
plants were examined.
Geranium crespitosum.-This plant is often rather abundant on the half-
gravel-slide, where its and give it importance.
Several plants were exammed, whICh ill general were sumlar (fig. 40). The
largest had a tap-root an inch ill diameter, which at a depth of 3 inches gave
off a strong lateral more than 1 cm. in diameter. This ran off horizontally,
following a curved course to a distance of 42 inches, and ended only 18 inches
below the surface. At a depth of 6 inches the root broke up into 3 parts, all
of which grew more or less horizontally, none of the branches reaching depths
of more than 37 inches. Most of these laterals approached the surface, the
profuse branches often ending within the second to the fourth inch of gravelly
soil and forming a fine system for surface absorption. Another lateral ran off
at a depth of about 15 inches to a distance of 52 inches from the base of the
plant. Geranium also has the same root habit of forming large mats of fine
rootlets, not only at the ends of the larger branches but also at the extremities
of the numerous smaller ones. The roots are reddish-brown in color, rather
brittle, and are uncovered with considerable difficulty. The older roots and
the crown are frequently more or less decayed.
Calamagrostis purpurascens.-This plant holds an important place in the
composition of the half-gravel-slide community. Like Elymus triticoides,
its root system is so well developed that it can compete successfully with most
other species (plate 26, B). Two clumps, each about 8 inches in diameter,
were examined. These were old plants with the flower-stalks 15 inches high
and in full bloom. From the base of the clumps almost countless numbers of
rather tough fibrous roots arose, the largest scarcely more than a millimeter
in diameter, while many were much smaller. Among them were many new
roots only a few centimeters long. These ran out in all directions like the
radii in a half sphere. The soil was well filled with those that extended verti-
cally and slightly obliquely to a depth of 18 or 20 inches, few of the longer
roots reaching depths of 32 inches, while the lateral extent of those running
vertically just beneath the surface was at least 2 feet on either side of the
plant. Many of the deepest and widest-spreading laterals maintain about
half their original diameter for a distance of 12 to 14 inches. From the very
base of the plant to the extreme tips the supply of fine rootlets is remarkable.
branch and rebranch again and again into conspicuous mats of almost
rootlets which penetrate every crevice of the gravelly soil. Few
If any of the grasses examined had such great masses of delicate clusters of
absorbing rootlets. Such a grass is excellently adapted to live in the half-
gravel-slide, as more thorough occupancy of the soil can scarcely be imagined.
In color the roots are brown to light tan. They were removed from the
gravelly soil with no great difficulty.
Kreleria cristata.-This grass, which is a characteristic dominant of the
is distinguished by an extremely fibrous root system. The
lOam roots at their outset vary from 0.2 to 0.3 mm. in diameter. At a depth
1 to 4 inches many of these run off parallel with the surface to a maximum
distance of 12 inches. These are branched and rebranched to the third
ra(nld' fourth order, the ultimate termini being almost microscopic in size
pate 26, A).
Gilla aggregata.-The long pink or red racemes of this abundant biennial
are very conspicuous on the half-gravel-slide, where the plant is quite abun-
96
r
dant. Of the 6 plants examined all had a large tap-root from 4 to 8 mm. in
diameter. The tap sends off numerous branches near the surface and tapers
so rapidly that at a depth of 10 inches it seldom exceeds 1 or 2 mm. in diameter.
The laterals are mostly wide-spreading, frequently nearly horizontal in posi-
tion, and much forked and rebranched into very abundant fine termini. They
reach distances of over a foot from the base of the plant. Like most plants of
this habitat the root system is not deep. No Gilia roots were found below the
28-inch level, but the moist surface soil is well filled with great quantities of
fine absorbing rootlets, especially to the depth of 18 inches.
Potentilla arguta. glandulosa.-This species frequently fonns families on the
half-gravel-slide. A large clump of these plants was examined. The com-
plexity of the rootstocks is such that they are hard to describe. The individual
clumps are connected throughout long distances by much-branched under-
ground parts, which often run horizontally or sometimes obliquely at depths
varying from 3 to more than 18 inches. These underground connecting parts
may reach a diameter of a centimeter, but are usually much smaller. They
are very much branched and give rise to the clusters of stems at intervals of
3 inches to more than a foot. At a depth of 2 to 5 inches the roots often throw
off a large number of small, short, but exceedingly well-branched surface
absorbing laterals. The larger roots are usually only 2 to 6 mm. in diameter
and run off in all directions, some to a maximum depth of 4.5 feet, while the
surface of the soil is filled with the extremely well-branched brush-like termini
of the laterals. These clusters are from 6 to 10 inches in length. The deeper
soils are also completely occupied by the irregularly branched and rebranched
root network, often to a depth of 4 feet (plate 26, c).
Frasera speeiosa.-This striking plant starts with a strong, rather fleshy,
glistening white tap-root, which is 1.5 to over 2 cm. in diameter. It penetrates
FIG. 41.-Root system of Fraaera apecWaa.
vertically downward to a depth of 9 inches, where it divides into 4 branches
of almost equal diameter. This main tap, throughout its length and even to
a short distance on the branches, was strongly marked by wrinkles, indicating a
considerable degree of contraction. Indeed, throughout the entire root system
not excluding the smaller branches, characteristic wrinkles and
occurred. At the point of origin the branches were about 7 or 8 mID. in diam-
eter. They traversed the soil in diverse directions, for the most part ruJ?Ding
more or less parallel with the surface, but giving off some vertically or
obliquely descending branches. The former sometimes ran off laterally to a
distance of 2 or 3 feet and at a depth of from 6 to 18 inches below the surface
of the soil. As the ultimate laterals approached their extremities they were
characterized by a marked, abrupt branching habit, which resulted in a tuft
of from 6 to 10 or more rather coarse, poorly branched termini varying from
2 to 5 cm. in length (fig. 41). The diameter of the laterals at the point where
they develop these tufts is often 2 mID. or more. Arising from these more or
less horizontal laterals, vertically descending roots were found to penetrate a
maximum depth of 24 to 30 inches. Like the tap, the branches of all orders
were more or less fleshy and very brittle, affording a decided capacity for food
storage. Several plants of different ages were examined. In general the above
description answers for all. C
Aster porteri.-This plant, which grows abundantly on the half-gravel-
slide, has the stem clusters connected by short rhizomes and possesses the root
system characteristic for plants of this habitat. From the base of the stems
arise a multitude of fibrous roots, the largest seldom being over 2 mID. in
diameter. Great numbers descend vertically or somewhat obliquely, some to
a maximum depth of 30 inches. Many others run off rather horizontally or
obliquely to a distance of 6 to 12 inches on either side of the plant before
turning downward. All are profusely branched and rebranched into So fine
absorbing network, so that every cubic inch of the soil for nearly So foot on
each side of the plant and to a depth of over 2 feet is forced to yield up its
available water and solutes to the excellent absorbing system.
HALF-GRA VEL-SLIDE ROOT SYSTEMS AND TIlE HALF-GRA VEL-SLIDE
ENVIRONMENT.
While the plants of the half-gravel-slide habitat still retain the
superficial and wide-spreading root habit of their predecessors of the
gravel-slide, this is supplemented by a much deeper portion which
extracts water and nutrients below the 18-inch level, as well as very
largely from the second and third foot of soil. The surface-spreading
roots may be explained by the frequent mountain showers which
supply water rather continuously to the shallow soils, while the
deeper-seated ones reflect the better deep-soil conditions, but especially
the more intense competition for water of this denser population.
The thick surface layer of loose rock fragments of the gravel-slide has <
here become more disintegrated and closely packed, and soon gives
way to soil. Hence water evaporates more freely. This, with the
competition of the taller half-gravel-slide plants, accounts for the
disappearance of gravel-slide species. The former shade those of
lesser height, but especially compete with them for water, undoubtedly
relying upon their deeper penetrating roots during periods of drought.
,VIII
5
Quadrat-bisect in the half-grave1 slide. The fsee of the trench was cut along the front of the
quadrat shown in Plate 26 A: S, Solidago ureop1&iltJ; A, Allium cemwm; E. ElvmUB ~
fragments of which are represented in blue; C, CalamagroBti8 ~ ; H, Beud&etv
parvifolia; B, BeBUt/a planlaginea.
PlATE 8
The soils of the half-gravel-slide are much more favorable for plant
growth. Although from one-third to one-half of the surface may still
be unoccupied, enough plants are present to prevent almost wholly
the slipping of the soil, even the thin surface layer of pebbles being
moved between plant clumps only during heavy showers. The first
8 to 10 inches of soil has a rich brown color, due to the presence of con-
siderable humus formed by the decayed vegetation. It has many more
fine particles and fewer large, coarse ones than the corresponding layer
on the gravel-slide. Although there is considerable variation, the rock
is decayed to a greater depth, due undoubtedly in part to the excre-
tions of plant roots and the resultant porosity of the soil, and to greater
water penetration following the death and decay of the roots. The
soil underlying the surface layer already described, while still con-
sisting largely of the decayed granite, is looser in texture and has
fewer large particles and much more sand intermixed with it, thus
affording a more congenial home for roots. In the gravel-slide only
local areas of soil about the sparsely spaced plants are filled with roots,
especially in the surface layer. In the half-gravel-slide, on account
of the greater number of plants and especially grasses, all of the soil
is well filled with roots to a depth of at least 18 to 20 inches or more,
while many roots penetrate to a depth of 3 feet.
The amount and distribution of the precipitation is practically
identical with that of the gravel-slide and the actual available water-
content is about the same also (table 18). This results from the greater
water-holding capacity of the soil, due to the presence of more humus
coupled with more perfect rock decomposition, and occurs in spite of
the increased absorbing and transpiring surface offered by the plant
population. The evaporating power of the air as shown in table 19 is
considerably less than that of the gravel-slide, owing to the greater
transpiration and shade and the reduced wind movement resulting
from the denser community. Notwithstanding the fact that one-
third to one-half of the soil surface may still be unoccupied, root
competition is rather severe. This is well illustrated in the bisect
shown in plate B, though this was made in a rather open portion of
the community. It shoUld be borne in mind that the roots seen repre-
sent only those actually occurring in a rectangle of soil 7 feet long and
4 inches wide. Undoubtedly competition is one of the large factors
in determining root distribution. This must be taken into account
with the soil conditions in reaching a logical explanation of the root
habits of the plants of this habitat.
VIII. TIlE FOREST COMMUNITY.
A forest community finally occupies the half-gravel-slide. It is
represented by Pinus ponderosa or Pseudotsuga mucronata, both of
which are frequently preceded by a chaparral stage. Along the
streams and moister slopes, Douglas fir meets the Engelmann spruce
Picea engelmanni, with which it often forms a mictium. A rather
number of herbs and undershrubs characteristic of the more mesophytie
type of forest were examained (plate 27, A, B).
Pirola chlorantha.-This evergreen herb is very abundant and forms
extensive clans on the floor of the spruce forest. The clusters of leaves arise at
intervals from a few inches to more than a foot from the glistening white
underground stems. These vary from 1 to 3 rom. in diameter, branch freely,
and form a connecting system for the individual plants. They lie at a depth
varying from 0.5 inch to about 5 inches. Just before the rootstock approaches
the surface to send up a cluster of leaves, it invariably branches, the branch
continuing to the next plant, etc. The root system is very meager and con-
sists of brownish roots arising at irregular intervals, usually about 1 inch apart
on the horizontal rootstock, although it is not unusual to find several inches of
the rootstock practically free from rootlets. These roots penetrate the moist
duff and rich humus soil to a depth of only 6 to 10 inches; while many of them
are only 1 to 1.5 inches long, others form brush-like clusters 3 to 5 inches in
length (fig. 42).
FIG. 42.-Pirola chlorantha.
Thalictrum fendleri.-This polydemic ranges in habitat from the halt
gravel-slide, into which it has worked its way from the
to dense spruce forest. In the Douglas fir forest it forms extensIve
The plants here described were examined in a spruce forest. The slend.
tops of these plants arise from a meager system of rhizomes almost black
color, from 0.5 to 2 rom. in diameter, and seldom over 6 to 8 inches long.
The root system springs from the rhizome near the base of the erect steIWI
sometimes as many as 30 or more fibrous roots originating from the base of
THE FOREST COMMUNITY.
101
'ngl
e
plant. For the most part the roots extend in a direction almost par-
with the surface of the soil. The depth at which they lie varies from
2 to 8 inches and the maximum lateral extent is 14 inches. Relatively a
'ery few of the roots take a more vertical or slightly oblique downward
but none reach a depth below 18 inches. The main roots are a milli-
meter 'or less in diameter. Throughout the course of the roots an elaborate
'stem of branching occurs. The branches ramify to the third and fourth
irder, and vary in .length from a few millimeters to 3 ?r.4 inches. their
extremities the mam roots are so well branched and dIvIded that the tIP con-
sists of a brush-like mat of fine rootlets (fig. 43).
FIG. 43.-Thalictrum fendleri.
Erigeron asper.-This plant fonns fine societies in the dense shade of the
spruce-fir forest. The individual clumps are connected by means of branched
surface rhizomes, seldom over and often less than 2 rom. in diameter, which
usually do not lie at a depth beyond 2 inches. From the base of the plant in
particular, as well as along the rhizome, a well-developed system of rather
fibrous roots arises. The largest of these do not exceed a millimeter in diameter
and are usually much smaller. While some of these penetrate to maximum
?epths of 26 to 30 inches, the first 18 inches of soil and especially the first 6
lDches are thoroughly filled with the hairlike and profusely branched rootlets.
Frequently these branches, which arise in great abundance all along the main
run off obliquely and even almost horizontally to distances of 5 to 10
lDches from the base of the plant (fig. 44).
f'. Erigeron macranthus.-This plant, which is not infrequent in openings in
propagates by means of rather coarse rhizomes often about 5 rom.
In diameter and several inches in length. From the base of the plant and from
the arise clusters of fibrous roots from 3 to 4 mm. to only 0.5
mm. m diameter. While many of these run rather vertically downward so
that some of the longer ones reach depths of 40 inches, others spread later:uty
to a of 14 to 18 inches and reach depths of only 4 to 12 inches. Thus
the soil for a of more a f?ot on either side of the plant and to a
depth of 2 or 3 feet li1 well supplied Wlth these fibrous roots. The branching
of .the I"O?t is almost with that of Erigeron asper (fig. 44), with which
this speCIes seems to mtergrade.
FiG. 45.-Fragaria virginiana.
FiG. .-Rhizome and roots of Erigeron FiG. 46.-AUium cernuum.
asper.
Fragaria virginiana.-Fragaria forms societies rather extensively on the
forest floor, even in the dense shade of spruce seedlings. In addition to the
long, slender stolons, it is furnished with rhizomes only 3 to 5 mm. in diameter
but often several inches long. From these arise the dark-colored fibrous roots
a millimeter or less in diameter. There are usually 6 to 10 of these on an inch
of the rhizome. Many of them pursue an oblique course to a lateral distance
of 4 to 10 inches from the base of the plant, where they reach a depth of 4 to
12 inches. Others penetrate more vertically downward to a maximum depth
of 12 to 14 inches. While the first inch of the roots below the superficial
rhizomes is rather destitute of branches, below this depth, especially in the
last 6 to 8 inches, the roots are supplied with an abundance of fine rebranched
termini ranging in length from 1 to 5 cm. (fig. 45).
THE FOREST COMMUNITY. 103
Allium cernuum.-This plant is very characteristic of half-gravel-slides, but
also occurs rather abundantly in the open portions of the fir forest. The plant
was examined in the forest habitat. The bulb, which is usually 12 to 15 mm.
in diameter, occurs at a depth of about 2 inches. From its base arises a cluster
of 10 to 20, or sometimes more, fibrous roots a millimeter or less in diameter.
These spread widely in the surface 6 or 8 inches of soil. Laterals were traced
to a horizontal distance of 8 to 12 inches from the base of the bulb, where they
ended at a depth of 4 to 8 inches. No roots were found at a greater depth
than 10 inches. These glistening white fibrous roots branch freely into
laterals from a few centimeters to 4 or 5 inches in length, but these secondary
roots are themselves scarcely at all branched (fig. 46).
Aralia nudicaulis.-This plant is often abundant on the lower slopes of the
fir and spruce forests, where it sometimes forms extensive communities. The
individuals are connected by a much-branched system of rhizomes, which
vary in diameter from 3 to 4 mm. to more than a centimeter. Frequently
they lie just below the soil surface, but they are quite abundant at all levels
and run in all directions and at all angles in the first 18 to 24 inches of soil.
FIG. 47.-Rhizomes and root system of Aralia nudieaulis.
As shown in figure 47, there arise from the nodes of the rhizomes rootlets with
clusters of well-branched termini, the whole usually not over 2 to 5 cm. long.
Often, however, rather well-branched roots 18 to 20 inches in length occur.
The direction in which these run depends somewhat upon the depth of the
rhizome. If the root arises from a rhizome near the surface it takes a vertical
or oblique course downward; at intermediate depths the roots frequently run
off horizontally, while from the deepest rhizomes it is not unusual to find them
ascending in an oblique direction toward the surface. As a whole, the root
system is rather poorly developed, probably due to the dense shade in which
the plant grows.
Opulaster opulifolius.-This shrub forms a very characteristic under-
growth in pine woods as well as in the open fir forest. Indeed, it reaches its
best development in the better light of the chaparral community and dies out
almost completely in forests with a closed canopy. Its r6le in succession is
not unlike that of Corylus (page 30). Two plants, 5 and 12 years old respectively
and about 3 feet high, were examined in the Douglas fir forest. From the base
of the clump prostrate stems about a centimeter or less in diameter run
horizontally just beneath the surface, or at least at a depth of only a few
inches, to distances of 2 or 3 feet, where they give rise to new plants. The
roots of this plant are often large and woody, sometimes a centimeter or more
in diameter, but usually smaller. They have a lateral spread of 5 to 8 feet,
but seldom penetrate the soil to a greater depth than 3 feet (plate 30, A).
From the base of the clump 6 to 12 of these roots arise. They run out laterally
in all directions, tapering gradually and branching freely and often dichoto-
mously, the ultimate branches being very minute and forming a great network
of absorbing brushes. From the main roots and their branches, as well as from
the rhizomes, great clusters of exceedingly well-branched laterals (from a few
centimeters to several inches in length) lay hold upon the moist soil of the
forest floor. In fact, the majority of the roots are confined to the surface 8
inches .. while only one or two branches were founq to reach the maximum depth
of 32 mches.
Ribes lacustre.-This plant is characteristic of rather dense moist woodland.
Four specimens, 6 or 7 years old and 1.5 to 2 feet high, were examined. They
spread through distances of 1 to 2 feet or more by means of rhizomes or
prostrate stems. From the base of the clump a tap-root 10 to 15 mm. in
diameter arises. Instead of descending vertically, this almost invariably
runs off in the surface soil, dividing rapidly into many major branches. While
many of these are only 2 to 4 mm. in diameter, they taper so slowly that at a
horizontal distance of 3 to 4 feet from the plant they may still be 1 to 2 mm.
wide. These long, tough surface laterals pursue their winding way through a
length of 6 to 8 feet, frequently reaching distances of from 5 to 7 feet from the
plant. They are usually unbranched or only poorly branched, save for
isolated brushes of small laterals, until within 12 to 18 inches from their tip.
Here they divide up into long, slender laterals all well-branched near their
extremities, but not at all to the same degree as Opulaster. While many of
these branches end in moist duff, others penetrate downward to a depth of
8 to 12 inches. The major portion of the root system is confined to the surface
soil, but a few branches were found at a depth of 2 to over 3 feet (plate 28, B).
Rosa. a.cicula.ris.-The rose frequently occurs in the spruce-fir forest, where
it is a relict of earlier stages in the succession. While under the lower light
intensity of the forest it is merely a low shrub, outside the forest it frequently
forms dense chaparral several feet in height. The plants are connected by an
extensive system of rhizomes which lie at a depth of 2 to 10 inches and vary
in diameter from 2 to 6 mm. These rhizomes run in all directions, branching
freely and at irregular intervals, giving rise to the stems which seldom reach a
height in the forest of more than 1.5 to 2.5 feet. The root system originating
from these rhizomes consists of both long and short roots. The shorter ones
vary in length from 1 to 12 inches. Almost from their point of origin they are
profusely branched and rebranched so as to form a brush-like mat. These
groups of thickly branched roots penetrate the surface soil in all directions,
forming a very efficient absorbing system. The longer roots vary in diameter
from 1 to 4 mm. at their point of origin and penetrate the soil downward in a
wandering course to a maximum depth of 2 to 5 feet. Throughout their course,
these longer roots are fairly well supplied with both short and long branches,
the shorter ones varying in length from 0.5 to 3 inches. The longer ones,
which are almost threadlike in appearance, may wander off in all directions,
even obliquely upward to a distance often of 2 feet or more. These branches
are rebranched much after the fashion of the main roots already described.
As the main roots and these larger branches approach their extremities, they
are divided several times so as to form fairly well-branched termini.
Arctostaphylos uva-ursi.-This ericad, which is the typical undershrub of
the pine forest, also occurs rather abundantly in the more open portions of the
Douglas fir forest, where it forms great mats covering areas of many square
meters. Its xerophytic qualities are shown by its frequent appearance in the
half-gravel-slide community. A very elaborate and well-developed root
system arises from the woody prostrate stems, which vary from a few milli-
meters to 3 cm. in diameter. While some of the roots reach a maximum depth
of from 40 to 46 inches, the major portion of the absorbing system lies in the
first 18 inches of soil. From the base of the prostrate stems as well as from
numerous shallow horizontal roots arise great numbers of rootlets only 1 to 2
mm. in diameter and from 4 to 8 inches long, but so abundantly supplied with
masses of branched and rebranched laterals that they fill the surface soil with
a network of roots. In addition to these, larger branches, sometimes even a .
centimeter in diameter, rwi off at all angles from the horizontal to vertically
downward, pursuing tortuous courses and branching and rebranching freely.
They attain a length of 3 or 4 feet and end at depths varying from a few
inches to more than 3 feet. The branches from these as well as the ends of
the main roots themselves form great mats of well-developed rootlets in the
crevices of the gravelly soil. Plate 28, A, shows something of the extent and
. abundance of these branches.
Senecio cernuus.-This herb is abundant in the fir forest. The root system
consists of a group of from 8 to 20 white, somewhat fleshy roots. At their
point of origin they vary from less than 0.5 mm. to 2 or 3 mm. in diameter.
Relatively few of these penetrate the soil in a vertical or slightly oblique
direction to a maximum distance of 12 to 15 inches. By far the greater part
of these roots take a course practically parallel with the surface at a depth of
from 1 to 3 inches-in fact, many of them run just below the surface accumu-
lation of duff and semi-decayed leaves; others may take a slightly more down-
ward course and end at a depth of from 4 to 8 inches below the surface. The
maximum spread of these horizontal roots may vary from a few inches to 18
or 25 inches. Throughout their course the main roots are fairly well supplied
with laterals varying in length from a centimeter or less to 6 or 8 inches.
These in turn may be branched again to the second and third order, so as to
form a very efficient surface system well adapted to absorb the shallow water-
content resulting from the frequent mountain showers. Eight plants were
examined (fig. 48).
Castilleia miniata.-This plant is a common component not only of grassy
half-gravel-slides, but it also occurs rather abundantly in the fir forest. Rela-
tive to the size of the plant it has a very meager root system. This consists
of 5 to 20 main branches, varying in size from 1 to 3 mm. These all show a
strong tendency to spread out laterally and run off in the surface soil, usually
at a depth not greater than 5 to 6 inches to distances of 12 to 18 inches or less.
No roots were found penetrating deeper than 8 inches. While these rather
coarse, yellow, brittle roots are fairly well branched, they are only poorly
supplied with fIDe absorbing .laterals as compared with other plants examined
in this community.
Heuchera parvifolia.-This plant is fairly abundant in the Douglas fir
forest. It has a strong tap-root, sometimes as large as 15 mm. in diameter.
However, this tapers so rapidly that within a distance of 6 to 8 inches it is
106 THE RELATIONS OF ROOTS.
usually not more than 1 to 2 IDm. wide (fig. 49). Of the 10 plants examined
only 3 had roots penetrating deeper than 6 inches. In these cases the tap
penetrated rather vertically downward, having no large branches but being
covered below a depth of 4 inches with multitudes of fine laterals only a.
centimeter or two in length. Near the tips they branch profusely and reach
depths of 15, 18, and 20 inches, respectively. Even in these plants the 2 or 3
FIG. 48.-8enecio cernUUB.
strong laterals, 4 to 7 IDm. in diameter, originated just below the crown and,
tapering rapidly like the tap-root, ran off horizontally at a depth of about 2
inches to a distance of 18 to 24 inches from the base of the plant. Unlike the
tap, these large laterals were well branched with rather widely spreading
rootlets, which gave off multitudes of tertiary branches, all of which ended in
a great network of absorbing rootlets. Many fine, much-branched rootlets,
only a few inches long, also originated from the crown of the plant where it
was buried in the moist duff. The root systems of those described were the
same as the others examined, except that the tap-roots of the latter group
took a course up the slope and assumed the r6le of laterals already described.
Saxifraga bronchialis.-This plant forms large mats, often several feet in
extent, which cover the moist soil on the forest floor or even occur on moist
rock surfaces. It is especially well developed in the Douglas fir community.
From the main root prostrate stems from 1 inch to over. 1 foot in length extend
out in all directions in the surface soil. These send up erect branches 5 inches
long the shorter ones being near the growing tip. As the soil accumulates
8 ; b o ~ t the bases of these erect stems, the lower leaves die and only an inch or
less of the living tip is exposed. These branches with their dead leaves and
accumulated debris occur in such density that they afford a congenial sub-
stratum for the abundant rootlets which arise from the horizontal portion
especially, as well as from the erect part of the stems. These delicate roots,
however, which vary from 1 to 14 inches in length, run off in a horizontal
direction in the moist soil just below the mat and seldom reach a depth greater
than 3 to 4 inches. They divide into fine, hairlike, well-branched termini and
occur in exceedingly great numbers. The main root or roots, depending upon
the size of the mat, are tough and somewhat woody, black in color, and from
2 to 3 mm. in diameter. They also run off horizontally and usually up tlie
slope, branching profusely and often dichotomously. They spread out some-
what fan-shaped and reach a distance of from 3 feet to over 5 feet from the
base of the plant. Although they were found to penetrate not deeper than
4: to 5 inches, their wide spread and the great numbers and extreme division
of their branches enable them to extract sufficient water from the surface soil.
Indeed, the termini of many of these branches consists of such a mat of deli-
cate, almost microscopic, ultimate branches that it is quite impossible to depict
them adequately either with the pen or the camera.
FIG. 49.-Heuchera parviJolia. FIG. 50.-Haplopappm parryi.
Haplopappus parryi.-This composite forms dense societies in the shady
woodland, the rosettes of long leaves almost completely covering the surface
over local areas (plate 27, B). It is supplied with a system of stout, much-
branched rhizomes that vary in diameter from 2 to 8 mm. and extend from
just beneath the surface to a depth of from 6 to 8 inches (fig. 50). From these
rhizomes arise large clusters of roots ranging in diameter from 2 mm. to less
than 0.5 mm. These roots descend rather vertically or somewhat obliquely
and completely fill the soil to a depth of 25 inches, while some reach a maxi-
mum depth of 28 inches. The younger roots are supplied with many branches
from their point of origin and are reenforced by a great network of branches
arising from the older roots in the deeper soil at 6 to 10 inches. The deeper
roots and their network of fine branches spread widely, often running for
several inches horizontally through the moist gravel soil. The whole root
system is surprisingly well developed, considering the habitat conditions
under which the plant grows.
FOREST ROOT SYSTEMS AND 'THE FOREST ENVIRONMENT.
A survey of the preceding data reveals the fact that the herbs and
shrubs of the forest floor are relatively shallow-rooted. Almost with-
out exception the bulk of the absorbing system lies within the surface'
18 inches of soil. Arctostaphylos and Rosa have roots which penetrate
deeper. The former, however, shows its xerophytic tendencies by
being the typical half-shrub of the open pine forest and often forming
large mats in the half-gravel-slide. In the Douglas fir forest it occurs
only in the most open places and disappears entirely in the spruce.
Rosa is a relict from an earlier stage of succession. It makes a much
better growth in full light, even in dry situations, where it forms dense
thickets. Even the roots of many of the Douglas fir and spruce trees,
including seedlings, saplings, and mature plants which were inciden-
tally encountered in the course of the work, were found to possess
many shallow roots (plate C). The root relations of forest dominants
- offer an exceedingly profitable field for investigation.
The water-content of the soil offers the logical exylanation for the
shallow root development of these forest plants. The forest soil is
usually deeper and always much richer in organic matter than that
of the half-gravel-slide. Usually there is a layer of 1 or 2 inches of
duff, beneath which the soil is very rich in decomposed humus to a
variable depth of 8 to 18 inches or more. Such a substratum furnishes
TABLE 20.-Average daily evaporating power of the air in the fore8t communitie8.
Date.
Douglas Engelmann
Date.
Douglas Engelmann
fir. spruce. fir. spruce.
c.c. c.c. c.c. c.c.
June 10-17 ......... 28.8 21.6 July 22-29 ......... 11.3 8.5
June 17-24 ......... 11.1 7.1 July 29 to Aug. 5 .. 16.0 11.4
June 24 to July 1. ... 20.4 15.0 Aug. 5-12 ......... 8.6 6.2
July 1- 8 ......... 15.2 11.1 Aug. 12-17 ......... 8.5 6.3
July &-15 ......... 5.7 4.4 Aug. 17-23 ......... 11.6 8.8
July 15-22 ......... 6.1 5.2
an excellent medium with a high water-holding capacity to catch the
precipitation of winter as well as that of the frequent summer showers.
The shade of the trees and various forest layers reduces the evaporat-
ing power of the air (table 20), while the water-loss from the soil is
further retarded by the layer of duff.
WEAVER
PLATEe
Pi F H F H F H H c
Quadrat-bisect showing root systems of shrubs and herbs of the forest floor. This was made along
the front edge of the quadrat shown in plate 27 B: H, Haplopappm parryij R, Rosa aciculariBj
Pi, Pinu8 fte:m1/isj F, Fragaria virginiana; C, Chamamerium angmtifoliumj P, Picea engelmanni.
That sufficient water ordinarily does not enter the soil to penetrate
deeply is shown by the following series of soil-moisture determinations
made during the summer of 1918:
TABLE 21.-WateT-content of the soil in the forest communities.
Depth of sample, Depth of sample, Depth of sample, Depth of sample,
Date.
o to 6 inches. 6 to 12 inches. 12 to 18 inches. 2 to 3 feet.
Fir. Spruce. Fir. Spruce. Fir. Spruce. Fir. Spruce.
June 10 ....... 6.6 12.7 3.1 8.1 ........ ........ ........ . .......
June 15 ....... 5.3 8.5 2.1 7.2 1.7 5.6 1.7 4.6
June 24 ....... 19.8 19.2 6.2 10.0 3.5 9.2 ........ ........
July 1. ...... 13.1 15.3 7.3 5.2 5.8 4.4 ........ . ........
July 8 ....... 8.1 12.8 5.4 5.2 4.8 8.4 ........ . .......
July 15 ....... 14.7 15.1 7.1 8.9 4.6 5.1 3.6 7.3
July 23 ....... 9.2 ........ 3.9 . ....... 2.7 ........ ........ . .......
July 29 ..... 16.4 18.8 11.3 9.2 8.6 5.8 ........ ........
Aug. 5 ....... 16.5 16.3 7.0 7.7 6.2 7.7 ........ ........
Aug. 12 ....... *18.0 10.3 *9.1 7.0 *6.7 4.3 *5.6 6.9
Aug. 19 ....... 16.4 21.0 7.8 11.2 3.2 8.2 ........ . .......
* Samples taken the day following a rain of 1.84 inches.
An examination of table 21 makes clear the fact that the greatest
amount of available water is in the surface 18 inches of soil. This
should account for the shallow root habit of forest herbs and shrubs.
The lesser extent of the root systems of these forest plants, when com-
pared as a group with the species of any of the preceding habitats, is
correlated with the more favorable aerial conditions for water con-
servation on the forest floor. Indeed, not oniy is the temperature
much lower, but the wind movement also is greatly checked. In the
:fir forest the average daily wind velocity was only 29 miles as com-
pared with 67 miles on the half-gravel-slide. Likewise, in the forest
diffuse light and low evaporating power of the air are the rule. Com-
pared, for example with the half-gravel-slide habitat, the aerial con-
ditions in the Douglas fir forest are only 54 per cent as severe. The
evaporation in the spruce forest is only 40 per cent as great as that on
the half-gravel-slide.
IX. ECADS.
During the course of these investigations, a number of species were
encountered and excavated in two or more different habitats. While
a few oftiherse have already been described in explaining the root
habits of plants of the several plant communities, others have been
reserved for comparison in this place. ~ Differences in ecads were
always determined by actual comparison of plant materials from the
two habitats at the same time, and these differences are expressed
wherever possible by means of photographs or drawings; Any personal
error was further checked out by the judgment of a second person,
no statements being made until there was a consensus of opinion . .r/
Smilacina stellata.-The shadefonn was excavated in a spruce-fir forest 24
feet from the brook bank and 3.5 feet above the water-level. The ground was
covered with about 0.5 inch of raw humus underlaid with about 2 inches of
well-decomposed humus mixed with sand. Below this was very moist sand
with gravel and rocks. The light value was only 0.02. .
The roots were much sparser and shorter, although somewhat greater in
diameter than those of the gravel-slide fonn (page 85). While an inch of the
rhizome of the latter fonn gave rise to 18 roots averaging 3.5 inches long, an
average inch of the shade rhizome showed 8 roots only about 1.5 inches in
length (plate 29, A). The longer roots, which have their origin near the base of
the vertical stems, were likewise both fewer and shorter. In the gravel-slide
form they varied from 2 to 5, in the shade fonn there was frequently none and
seldom more than 2. The maximum length of the latter was only 16 inches,
with an average length of about 11 inches; in the former case roots were
traced to 44 inches in depth, with an average depth of about 32 inches. An
intermediate stage was found in the fir forest, a habitat which has been shown
to be somewhat less meosphytic than the spruce. The roots were nearly as
sparse as those in the spruce forest but somewhat longer.
Chamrenerium angustifolium.-This cosmopolitan plant is found in habitats
of all degrees of mesophytism, from the gravel-slide and bare burn to very
moist dark places in the spruce forest. A large group of these plants was
examined at the foot of a slope near a stream and 4 or 5 feet above water-
level. They were growing in the half shade of Douglas fir and mountain
maple on one side. Below the first inch or two of duff and humus the soil
consisted of a rich black sand intermixed with gravel and rich in humus to a
depth of 2 or 3 feet. The plants are connected by means of strong, tough,
well-branched rhizomes from 5 mm. to more than 10 mm. in diameter. These
lie at depths varying from only 0.5 inch to 6 or 8 inches. The subterranean
portions of the erect stems are practially free from roots. At rather distant
intervals along the rhizomes, roots varying from 1 to 5 mm. in diameter
arise. Many of these pursue a more or less vertically downward course, with
but a few major branches, and reach depths of 35 to 48 inches (fig. 51). They
taper very gradually and are only fairly well clothed with small rootlets.
The latter usually occur in clusters of from 3 to 7 and are frequently less than
~ inch long. In addition to these, however, other longer and abundantly
branched laterals supplement the absorbing system. These extend from 3
to 10 inches. Sometimes these deep roots break up into numerous branches,
all of which are similar to those already described. In addition to these larger
roots, the rootstock also gives rise to smaller, shallower, and frequently
ECADS. 111
obI" que laterals. In fact, the larger roots also sometimes take an oblique
course. As indicated in the figure, the root system is somewhat meager. The
older roots may be identified by their brownish-yellow and somewhat flaky
cortex and the tough, rope-like stele within, while the younger roots can be
told by the clustered arrangement of their branches.
FIG. S1.-Root systems of ecads of Chamrmerium an{J'U8tifolium: a,' b, forest.
A number of plants were examined on a steep gravel-slide. Like those
in the fir forest, they were mature and in full bloom. Those on the gravel- .
slide ranged from 1.5 to 2 feet in height, while the shade forms were 4 to 5
feet tall. The most noticeable difierences in the underground parts were
the shallowness and linear extent of the root systems and the remarkable
difierence in number and fineness of the smaller roots. Of the several plants
examined on the gravel-slide, all sent roots up the slope at a depth of 2 to
10 inches or more. These frequently ran rather parallel with the surface,
often branching into major branches, for a distance of 3 to 6 feet or more.
From these rather shallow roots some of their branches pursued an obliquely
downward course. None of them reached a depth greater than 20 inches,
but almost invariably turned and ran parallel with the surface of the soil.
A comparison of the two root systems emphasized the paucity of small
laterals in the forest ecad and their abundance in the gravel-slide. In the
latter habitat, great clusters of fine rootlets (resulting from the repeated
branching of the main roots, especially near their extremities) filled the soil
with a web-like network in marked contrast to the rather sparse, absorbing
rootlets of the shade form.
Elymus triticoides.-A family of Elymus triticoides, which formed a sod
completely covering the ground under a canopy of Douglas fir, was examined.
The soil was fairly well decomposed and filled with humus to a depth of 18
inches. The underlying soil was very similar to the second foot of soil in the
half-gravel-slide. A trench 4 feet long was dug through the midst of the com-
munity and careful examination showed that no roots penetrated to a depth
greater than 38 inches, while in the half-gravel-slide a maximum penetration
of 46 inches by the larger roots was found (page 92). A marked difference was
, evident both in the density and coarseness of roots, especially in the first 2
. feet of soil. In the half-gravel-slide the roots were not only much coarser, but
they also filled the soil much more completely. In the shade form the flowering
stalks were also much less abundant. The ecads of this species show differ-
. ences in depth of penetration and in the number of roots per given area, as
i well as in size.
Bouteloua gracilis.-This widely distributed and important grass has been
studied under four different habitat conditions. In the prairies of eastern
Nebraska it occurs often in fairly pure stands on the lighter soils of gravelly
ridges and is also frequently found dominating areas of alluvial soil on bottom
lands. One of the several plants obtained from a gravelly ridge at Belmont
is shown in plate 6, B, while the root system is fully described on page 8. A
lew roots reached a maximum depth of 46 inches, although below 2 feet they
were very sparse. Great masses of roots occupied every cublic centimeter of
the soil to a depth of 18 inches. Other groups of plants examined in two
locations on alluvial soil showed a somewhat poorer development of the root
system, but the general distribution and depth were very similar to those
growing in the gravelly soil.
This grass was again examined on the plains of Colorado, where Bouteloua
is the dominant, but where societies of Ari8tida purpurea, Psoralea tenuiftora,
and Artemisia frigida, with other deep-rooted plants, indicate rather favorable
water-content conditions. Here the soil was found to be well filled with these
rootlets to a depth of 30 inches, while in the next 6 inches they were still fairly
abundant, some of the longer ones penetrating to a maximum depth of 48
inches. The lateral spread in the surface soil was much more pronounced than
in the plants examined near Lincoln. The plains species have many wide-
spreading and exceedingly well-branched roots filling the surface soil and
extending to 1.5 feet or more on all sides of the clump.
This grass was likewise studied in the typical hard-land or short-grass
country about 25 miles southeast of Colorado Springs (p. 65, plate 20, A),
where Bouteloua again dominates and the shallow-rooted Opuntia polycantha
is the principal species, a community conspicuous for the almost entire
absence of Psoralea, Gutierrezia, Chrysopm, and other deep-rooted dicotyle-
dons. A number of roots were found to penetrate to a depth of 51 inches,
while they were very abundant to a depth of 40 inches. Their wide-spreading
habit and their position in the soil was almost identical with those in the plains
habitat just described.
Finally, a fourth group of these plants was excavated in the sandhills
(p. 68), about 40 miles southeast of Colorado Springs but under the same
general condition of rainfall. A trench 8 feet long and 5 feet deep was dug on
a well-covered sandhill where this species was the dominant and formed
rather pure stands. From the short, tough rhizomes, usUally only an inch or
ECADS. 113
two in length, great numbers of tough, fibrous roots a millimeter or less in
diameter arise. On 4 cm. of one of these rhizomes 53 roots originated. While
some of these pursue a rather vertically downward course to a maximum
depth of 27 inches, or wander out somewhat obliquely, great numbers, perhaps
half, run off more or less horizontally with the surface of the soil, reaching
distances varying from 10 to 25 inches on either side of the clump (plate 29, B).
Some of these ended only 2 to 4 inches below the soil surface, the tips of the
growing roots being characterized for a distance of 1 to 3 inches by the entire
absence of lateral branches. In fact, some of these had penetrated so near the
surface that the soil had dried out and the root-tips had died. However; on
mature roots, not only are the tips but also the entire root system densely
clothed with great masses of capillary, much-branched rootlets of varying.
lengths, exclusive of the first 2 to 4 cm., which lie in the dry sand. Most of
these branches are only 1 to 3 cm. in length, but frequently the main fibrous
roots, especially the deeper-lying ones, repeatedly give off laterals many
inches long, which run off at various angles from the course of the main root.
While the first 8 inches of soil are literally filled with great masses of this
wonderfully efficient absorbing system, the lower soil stratum to 2 feet is also
abundantly supplied with fine roots.
The marked difference in the root penetration of the sandhill ecad as con-
trasted with those of the plains may be explained upon the basis of a fairly
abundant water-supply in the shallower soils (first 2 feet) of the sandhills as
contrasted with the uniformly drier soils of the plains. As pointed out on
page 79, the surface sand forms a splendid mulch which inhibits water-loss by
evaporation. It may be noted that the plants from the gravelly soils of the
knoll in the prairie near Lincoln are intermediate in root depth between the
sandhills and the plains ecads.
Stipa comata.-As described on page 53 and illustrated in plate 19, A, this
species is deep-seated in the hard, dry plains soil. The surface 28 inches of
soil is thoroughly occupied by the main roots, which are only a few millimeters
apart, the interstices being completely filled with horizontal branches. Even
to a depth of 32 inches the soil is fairly well filled with much kinked and
rebranched threadlike rootlets, while not a few reach a maximum depth of
over 5 feet. In addition, the lateral spread of the shallower oblique roots is
18 inches or more on either side of the base of the clump. The sandhill ecad
of this species, while having a similar surface-root distribution, stands out in
marked contrast as regards its superficial position in the soil, a condition
already explained upon the basis of a relatively high water-content of the
shallow soil.
Tufts of Stipa comata about an inch in diameter were abundant on the slopes
and tops of captured sandhills. Of the 5 clumps examined, rather large r ~ o t s
(to the number of 15 to 25) originated from the base of the crown. None of
these exceeded a millimeter in diameter. While some of them descended rather
vertically to somewhat obliquely, a few reaching a maximum depth of 23
inches, the remainder ran off much more obliquely, having a lateral spread of
10 to 15 inches and reaching depths varying from 3 to 20 inches. Although
the roots are not branched in the first 1 or 2 inches of soil, they are densely
covered with hairs. Upon entering the moist soil they immediately begin to
branch, giving off countless numbers of very fine, mostly rebranched short
laterals only a few millimeters to an inch in length. At a distance of from 8
to 12 inches from the tip, the main roots frequently break up into 3 to 5
laterals several inches in length. These spread out in all directions and their
profusely branched and rebranched network of finer absorbing rootlets lays
hold upon considerable areas of soil. .
Yucca glauca.-The root systems of numerous individuals of this species
were examined in the plains soil (page 51) and also in the half-gravel-slide of
the mountains. The main roots, which are more abundant in th!Ol half-gravel,
slide ecad, are much more profusely branched. Great clusters of long, well-
branched sublaterals fill the soil iI! sharp contrast to the poorly branched
laterals characteristic of the half-dozen yuccas examined on the plains (fig.
52). A marked difference was also noted in the diameter and the fleshiness
of the system of roots. While most of those of the plains were rather dry and
often papery, those of the gravel-slide were generally more turgid, thicker,
and fleshy. The rhizome system is somewhat more branched and is nearer
the surface than on the plains.
I ~
if' < ~
inch
,
~ a ~
FIG. 52.-Fragments of the roots of Yucca glauca: a, plains form; b, half-gravel-slide form.
Allionia linearis.-This four-o'clock is a widely scattered secondary species
on the plains and also occurs rather abundantly in the sandhills. Of the 8 or
more plants examined in the former situation, all had strong and rather
fleshy tap-roots, the largest 12 mm. in diameter. However, the tap is not
uniform, often contracting at the top and being largest at a depth of from 8 to
16 inches. At about 2 feet in depth it narrows down abruptly to scarcely more
than 2 mm. Except for small curves and kinks, it pursues a vertically down-
ward course, some to a maximum distance ef 5.5 feet. The first 18 to 24
inches of the root are practically unbranched, and in fact the lower part of the .
root system has no long laterals. Those that do occur at irregular intervals
consist of groups of hairlike laterals, seldom more than a centimeter in length
and usually only 4 or 5 mm. long. These are very minute in diameter and
rather well branched. The tip ends abruptly with only a few short branches.
The root is dark brown in color and very brittle, the interior parts being
glistening white. At a depth varying from 6 to 14 inches, usually one or
sometimes two laterals arise. These may be as large as 2 mm. and run off
horizontally or slightly upward, ending 18 inches to more than 2 feet from
the base of the plant. Like the main root, they are only poorly clothed with
clusters of short branches. (fig. 53).
Other specimens were examined in the sandhills from a half-captured
blowout. Five roots were examined. In general they were very similar, the
strong taps (which were about 5 to 8 mm. in diameter) descended rather
vertically to a depth of 4 or 5 feet. A typical specimen is shown in figure 54.
One to three branches, usually only one, occurred in the first foot of soil. The
lower portions of the roots had a few branches or were in some cases almost
destitute of them. The tops were both greater in diameter and taller than the
plains specimens examined. The leaf surface also was greater.
ECADS.
115
From the above, as well as from figures 53 and 54, it may readily be seen
t h a ~ Allionia lineam is a rather stable species as to root system, the root habit
differing only slightly in the two habitats. The clusters of small branches
were quite pronounced in the plains ecad, while this was compensated for in
the sandhill form by a somewhat greater degree of branching. The habit of
sending off one or more large and rather horizontal branches in the surface
foot of soil was characteristic in both habitats, as was also the strong tap-
root, which showed ,bout the same degree of penetration in both groups.
--....
1
~
--:r
!
~
I
)
..
,.
\

... ~
rh
FIG. 53.-AUionia linearis from the (,.. FIG. M.-AUionia linearis from the
plains. sandhills.
~ - " "
Abronia fragrans.-This plant is rather widely distributed over the plains
in disturbed areas, where, as in the sandhills, it frequently forms socies. The
large, showy, hemispherical clusters of flowers make it very conspicuous.
Three plants were examined. The description for one will answer in general
for all. The two largest had tap-roots 6 and 7 rom. in diameter respectively;
the smaller was only 3 rom. in diameter. Both of the larger plants had 3
stems which reached a height of 14 to 16 inches and were either in blossom or
in fruit; hence these may be considered typical mature plants. The tap-roots,
except where they twist and curve irregularly back and forth for an inch or
more in crevices of the cloddy soil, pursue a vertically downward course. The
smallest reached a depth of 34 inches, the largest 40 to 43 inches respectively.
As a whole the root is very poorly branched, no large laterals being given off.
Beginning near the surface, however, and extending quite to the tip, short
threadlike laterals, seldom exceeding 0.2 to 0.3 rom. in diameter and often
FIG. 55.-Abronia fragra'TUl from the sandhills.
arising in groups of 2 or 3, rim out laterally for a distance of less than 1 cm.
to more than 4 inches. The longer ones occur within the surface foot of soil.
These branches are very scattering, often 3 or 4 cm. of root having no branches
whatsoever. The tip is only poorly branched. The older roots are black in
color, this being confined to the outer layer of cortex, the interior being white.
The younger roots vary in color from brown to yellowish. They are removed
only with extreme difficulty from the hard soil. The tiny lateral rootlets are
seldom branched beyond the second order and do not form a network (plate
29, c).
Four more rather large plants, all in flower or fruit, were examined. They
were growing on a captured sand-dune and had tap-roots varying from 6 to
10 rom. in diameter. One of the largest of these is shown in figure 55. The
tap is prominent throughout and tapers slowly to the tip, which reaches a
depth of from 40 to 48 inches. The plant is characterized by 2 types of
branches; the shorter ones vary from 2 to 20 rom. in length and occur in tufts
ECADS. 117
or clusters, both on the main root and larger branches. Besides these, all
of the roots examined had 3 to 6 or more larger branches, which ran off from
the tap somewhat horizontally at all levels and to distances varying from a
foot to over 3 feet. As shown in figure 55, the root system as a whole
is poorly branched, although the sandhill ecads were much better provided
with laterals and smaller rootlets than were the specimens examined in the
plains soil. The root is yellowish-brown to nearly black in color. It is very
brittle, the laterals especially being followed with extreme difficulty.
A comparison of these descriptions, together with an examination of the
two figures, reveals the fact that the sandhill form, while showing the same
habit of a strong tap-root which penetrates to about the same depth as the
plains species, differs in its more abundant branching as well as in the great
lateral spread of the branches. In these respects the modifications are not
unlike those of other sandhill ecads already described.
Kreleria cristata.-This very important and cosmopolitan grass was first
examined in the prairies of eastern Washington, where it is a common bunch-
grass. Under the prevailing conditions of rather moderate precipitation
(about 21 inches), most of which falls in winter and is followed by a season of
summer drought, it flowers in late June, soon dries up, and remains dormant
until revived by the autumn rains. In this region the deepest root found was
at 28 inches and 15 inches was determined as the average maximum depth.
In the prairies of eastern Nebraska it is also an important grass, growing under
an annual precipitation of 28 inches, much of which falls during the growing
season. Here none of the roots of the 7 plants examined reached depths of
over 21 inches, while the average maximum root depth was about 15 inches.
Kmleria was further examined in the hard plains soils of east-central
Colorado and under an annual (summer) precipitation of about 15 inches.
Here the root distribution was almost identical in lateral spread and depth
with those already described. The greatest depth was 26 inches and the
average maximum depth about 14 inches. Finally, the same species was
excavated on a half-gravel-slide at an elevation of 9,000 feet in the Rocky
Mountains. As was characteristic in the other habitats, the first 6 to 8 inches
of soil was completely filled with the profusely branched and matted root
system. Roots were quite abundant to a depth of 12 inches, while some of
the longest penetrated to 18 or 20 inches. .
A comparison of plates 5, A, and 26, A, showing the roots of plants from the
prairies of eastern Nebraska and from the gravel-slide respectively, reveals
the fact that they are almost identical. Compared with the profound differ-
ences in the ecads of Stipa comata and Elymus triticoides, Kmleria is a remark-
ably stable species.
Chrysopsis villosa.-This composite is widely distributed throughout much
of the grassland formation, occurring abundantly in the sandhills (plate 21, A).
On the plains several plants were examined. All had strong tap-roots, which
were woody for 2 or 3 feet. The height of the plant above ground was 6 to 10
inches. A second plant had a tap-root with a diameter of 12 mm. It pene-
trated almost vertically downward, except for local curves and kinks, usually
less than an inch from a vertical line, to a depth of 8.6 feet (fig. 56). The first
18 inches of the tap, beginning about an inch below the soil surface, is abun-
dantly supplied with threadlike laterals, mostly less than a millimeter in
diameter, which run out in a horizontal direction from a distance of 1 inch to
more than a foot. These laterals are practically free from branches. At a
depth of 6 to 18 inches, half a dozen stronger laterals, each about 1 mm. in
118 TIIE ECOLOGICAL RELATIONS OF ROOTS.
diameter, ran off rather horizontally for a distance of 18 inches or so before
turning downward. At 2 feet the root had a diameter of only 2 mm. and was
very much twisted and curved, as is characteristic of many plants in pene-
trating the hard soil. Practically no branches were given off to the very tip,
which was threadlike and unbranched. At a depth of 6 feet, where the soil
became looser and more moist, the roots pursued an even course, in striking
contrast to the twisted parts in the harder soil. The younger portion of the
root is almost white in color and covered with a dense coat of root-hairs. The
older parts are dark brown. This description answers for two other plants
which were growing near it.
Another plant gave off two laterals at a depth of 18 and 21 inches respec-
tively. These were each a millimeter in diameter and ran rather horizontally
to a distance of 2 and 2.5 feet. At a depth of 34 inches another lateral, 2 mm.
in diameter, came off and ran almost parallel with the tap. This plant had a
woody tap 3 cm. in diameter, which split at the top and gave rise to three
clusters of stems. It tapered rapidly, however, and at a depth of 2 feet was only
4 mm. in diameter. The main root outside of branches already mentioned
branched but little, as indicated in the drawing. It reached a depth of 13 feet.
On a half-captured dune in the sandhills, 5 or 6 large mature plants were
examined, all of which agreed in having a multicipital crown arising from
the 2 to 5 upper forks of the thick, woody tap-root. The surface branching
was exceedingly well developed. From the very base of the plant numerous
lateral branches arose, varying from 0.5 mm. to 60r 8 mm. in diameter. All
. were fairly well rebranched, the smaller laterals and the larger branches of
the major ones being densely clothed with root-hairs. In most cases the
tap-root, owing to the enormous numbers of large lateral branches, rapidly
diminished in size, so that at a depth of 18 inches none were larger than 5
mm. in diameter. This stands out in marked contrast to the stout, deep-
seated tap of the plains. Indeed, the deepest root examined penetrated to a
depth of only 6 feet 11 inches, while most of them reached depths of only
4 to 6 feet. The lateral branches, both large and small, formed a dense
network of roots filling the soil from 18 to 24 inches on either side of the
plant. All of the larger roots were well supplied with small, well-branched
absorbing laterals. The divisions of the tap-root were themselves branched
and rebranched to such an extent that, as they passed downward, great
numbers of roots were found penetrating the soil to depths of 4 or 5 feet.
Still other plants were examined which confirm the above descriptions,
the taps and larger branches having a strong tendency to run obliquely or
even horizontally, rather than vertically downward as in the plains ecads
(plate 30, B). .
This affords a very clear case of the effect of the habitat upon root develop-
ment. The plains form has a root which is approximately twice as deep-
seated as those in the sandhills. While the tap is prominent throughout in
the former, it soon loses its dominance in the sandhill form and often scarcely
exceeds in importance some of the stronger laterals. While both forms are
supplied with rather abundant surface laterals, in the plains form these are
largely confined to the surface foot, while in the sandhills they occur to a
much greater depth and are abundant along all of the major branches. The
branches in the first foot of the plains soils were. mostly less than 1 mm. in
diameter and had a horizontal spread seldom exceeding 18 inches. They
were practically free from branches. In the sandhills the laterals ranged
from 0.5 to 8 mm. in diameter and were rather well branched. In number
these lateral branches, both large and small, formed a dense network of roots
ECADS. 119
fillin the soil to a distance of 18 to 24 inches on either side of the plant.
Such
g
marked differences in root habit are probably accompanied by marked
changes in the aerial parts. This is a rich field for future investigation.
Euphorbia montana.-This spurge is rather widely distributed throughout
the plains and mountains and is often locally . abundant. :rhose . examined
on the plains had a strongly marked tap-root, which started WIth a diameter of
6 mID. and pursued a tortuous and zigzag course dmrnward to a depth of
75 feet where it ended in pearly white, hairlike branches (fig. 57). Within the
fust there were 4 short branches from 4 to 8 inehes long, which were
themselves slightly branched. The larger branches from this point downward
were very sparse, there being only one of note, which eame off at a depth of
about 20 inches and pursued an obliquely downward eourse for a distance of
about 2.5 feet. Along the whole course of the root below the first 21 inches,
very fine, pearly-white, hairlike branches were found in abundance. Par-
ticularly was this true from 21 to 55 inches deep. In this region a large soil
fissure occurred, and these pearly-white branches, oftentimes arising in pairs,
were matted and profusely branched, extending horUontally from the main
root to a distance of 10 inches in some cases. The matting of these small
branches in the fissure was probably due to the excess of water which came
down from the surface. Below this fissure these hr.mehes continued, but
much fewer in number, to a depth of 60 inches, where a very moist, easily
penetrable stratum of soil was encountered. In this stratum the small
branches again occurred in great numbers, being sometimes 6 to 8 inches long
and as many as 25 per linear foot. Here, too, they often occurred in pairs.
The root is light brown in color. It has a fleshy corte:s: and a tough,fibrous
stele; when broken a very small amount of latex exuded..
Compared with this root type, that of the half-graYel-slide, with its shal-
lower but more widely spreading and much more profa!iely branched roots,
stands out in marked contrast. It shows a profound series of modifications in
adapting itself to this very different habitat. This is evident from a com-
parison of figures 57 and 58, notwithstanding the fad that the half-gravel-
slide plant here illustrated was exceptionally deep-rooted, and from the follow-
ing description, which is typical of one of several other half-gravel-slide
plants examined.
This was a large plant, with a tap an inch in diameter, which divided at a
depth of 1 inch into two nearly equal parts. The one r.w up the slope in a
horizontal position and at a depth of only 4 to 10 inches. It soon broke up
into many smaller branches, which diverged in all but.a downward direction
and extended to distances of 3 to 4 feet, and mostly only a. few inches below
the soil surface. The second descended vertically, but p,ve off 2 branches 5
and 8 mm. in diameter, respectively, at a depth of 2 and 3 inches, while the
tap was now only 5 mm. in diameter. The largest of these branches ran off
at a depth of only 8 inches to a lateral distance of 5 feet. At 16 inches from
the base of the plant it broke up into numerous equal-9..md branches which
rebranched profusely. None of these reached a depth greater than 14 inches,
while many of the obliquely upward running branches fiDed the soil to near
the surface with great masses of finely branched rootlet&. The tap tapered
very gradually and after branching considerably, JIlO5tly with vertically
or oblique or even horizontal branches. reaebed a depth of 58
mches. The spread of the deeper branches on either side of the tap was only
about 12 inches. Tiny, fragile, glistening-white mala of branches form a
network of rootlets 2 to 8 inches long, surrounding all but the larger parts of
the older laterals. Four plants were examined.
Flo. 57.-Euphorbia montaM from
the plains.
FIG. 58.-Euphorbia montaM from the
half-gravel-slide.
ECADS.
ROOT MODIFICATIONS OF PQLYDEMICS .
. Of the 10 polydemic each growing in at least
two different habitats, 7 show very striking changes in their root habits,
2 made practically no change, while 1 exhibited only moderate differ-
ences of root development.
Bouteloua, Stipa, and Chrysopsis growing in the plains soil were all
deep-rooted species, the latter indeed reaching a depth of over 13 feet.
In the sandhills they all conformed to the typical root habit of most
plants of this habitat. None of them reached more than half their
former depth, but the shallow lateral root development and lateral
spread were much more pronounced, especially in Chrysopsis. On the
other hand, the more stable species as regards root development;
Allionia linearis and Abronia jragrans, when growing in these same
habitats, were only slightly or not at all modified. The former showed
practically no difference in the sandhill and plains ecads; the latter
had a tendency to develop more and longer branches in the sand.
These, with Kreleria, which was examined under four distinct sets of
conditions in widely separated areas and found to be practically the
same in all, form a very conservative group as regards root type,
although, as shown by E. Clements (1905), the aerial parts of some are
very plastic. This forms a unique problem for further investigation.
Euphorbia and Y mea, while conforming to the plains root habit
in having a rather deep and widely spreading absorbing system, were
found to be profoundly modified when grown in the half-gravel. In
the former the depth. of penetration was always much less, while the
branching, like that of y' ucca, was much more pronounced.
Of the three species, Smilacina stellata, Elymus triticoides, and
Chamamerium angustijolium, examined in gravel-slide or half-gravel-
slide and in the forest, all took on the typical root habit characteristic
of the several habitats respectively. All had fewer branches, a smaller
lateral spread, and a lesser depth of penetration in the forest than in the
open. The rather great root depth of Chamrenerium in the Douglas
fir forest is an exception. This is somewhat puzzling and seems to be
quite contrary to the usual rule of root development.
X. SUMMARY.
This study was undertaken to determine the root habits of dOminant
and subdominant plants growing under a wide range of climatic and
edaphic conditions; to find the root relations of the plant communities
as units of vegetation; and to determine the root distribution and root
competition of the individual species in their relation to other speCies
in the community. Other aims were to determine the relation between
the root habits of plants in various communities and their successionAl
sequence, and to obtain a more definite knowledge of the indicator '
value and the significance of various species used in classifying lands
for grazing or for agriculture, as well as to aid the forester in selecting
sites for afforestation or reforestation. . ' .... ..
The investigation extended over four years, during which tinie
more than 1,150 individual root systems of about 140 species of
grasses, and other herbs were excavated and studied. These species
were members of 8 different plant communities widely separated
graphically and growing under distinctly different climatic or edaphic
conditions. The commlID,ities studied were the prairies of eastern
Nebraska, the chaparral'Of southeastern Nebraska, prairies of south-
eastern Washington, the plains association and sandhill subc1inUn of
Colorado, and the gravel-slide, haH-gravel-slide, and forest communitief
_ of the Rocky Mo.untains of Colorado. .' '. "; '" ',.
The trench or pit method was used in excavating the roots, the roof
system being removed with appropriate apparatus from the face of the
trench. In case of roots extending to depths beyond 15 feet, tWQ
working levels were maintained. The quadrat-bisect method has been
employed to show the root systems of communities in. place and thus
exhibit their interrelations in detail. ' '7>."
The rOQt systems of 33 species were examined in the prairies: of
eastern Nebraska; 13 of these were grasses. All of the dicotyledonous
plants were found to extend well beyond a depth of 2 feet, while the
roots of 6 of the grasses were confined to the surface 2 feet of soil.
Four of the grasses and 5 other species were found to penetrate well
below the second foot of" soil, but seldom deeper than 5 feet. More
than haH of the plants studied, including three dominant grasses, have
roots which reach depths greater than 5 feet-indeed, most of them.
penetrate to distances of from 7 to 9 feet and a few to a maximum
depth of from 13 to 20 feet.
These prairie species grow under a mean annual precipitatioI
of about 28 inches, over 20 inches of which fall during the growing
season. The soil is of the type commonly called loess, much of which,
however, is confounded with glacial drift. Soil-moisture determina-
tions for more than two seasons show that the water-content is some-
times reduced to such a degree that no moisture to a depth of 4 or 1
SUMMARY. 123
feet is available for plant growth. The wind, prevailingly from the
south or southeast during the growing season, averages about 70
miles per day over the vegetation. The mean summer temperatures
are high (70
0
to 75 F.), while the mean humidity is often low (58 to 70
per cent). The average daily evaporation throughout the growing
season is about 21 c.c.
The plants of the prairie community, in response to these environ-
mental conditions, have developed very efficient widely spreading and
deeply penetrating root The l?rairie root habit
emphasizes depth of penetratlOn and WIdely spreading, deep laterals
much more than the desert community (Cannon, 1911, 1913; Markle,
1917). The roots of prairie plants are grouped into more or less definite
absorbing layers, many of the deeper-rooted species having few or
no absorbing roots in the first few feet of soil. The layering of the
roots reduces competition and permits the growth of a larger number
of species. ..
The root systems and mechanism of invasion of several dominant
shrubs of the chaparral community, which occupies the tension zone
between forest and prairie, were examined. All are supplied with
splendid absorbing systems which are somewhat variable as to depth
(5.5 to 21 feet), but.all of which are deep-seated. In addition, they
all have excellent methods of vegetative propagation. Their presence
indicates a soil richer in humus and higher in water-content than
ihat of adjacent grassland areas. These differences, as well as a lower
evaporating power of the air, are brought about by the reactions of
the shrubs. These consist of the accumulation of wind-blown snow and
plant debris, a lowering of. the light values and a consequent disap-
pearance of most or all of the prairie species, obstruction to wind
movement, lower temperatures and higher humidity, in part due to
shade, and a less rapid oxidation of the humus materials. ..
The root systems of 18 dominant and subdominant species were
examined in the prairies of southeastern WaShington. These prairies
represent an extreme westward extension of the great grassland
formation lying east of the Rocky Mountains. They are characterized
by the absence of late-maturing grasses and in general by an earlier
seasonal development. Three of the four dominant grasses have root
systems confined to the surface 18 inches of soil. This is correlated
with early seasonal growth and maturity. The fourth, like the other
herbaceous species, has a deep, widely spreading root system. All of
these reached depths from 4 to 6 feet, while some penetrated even
deeper. However, the plants of the prairie community are not so
deeply rooted as those of the prairies of eastern Nebraska. This may
be correlated with the environment. .
These prairie species grow under a mean annual precipitation of
21 inches, only about 7 inches of which fall during the growing season.
But the silt-loam soil is extremely retentive of water, so that the
precipitation, which occurs so gently that there is little run-off, is held
as in a .great reservoir that is rather thoroughly emptied during the
following summer. Soil-moisture determinations for two seasons
show that the water-content is sometimes reduced to the non-available
point for plant growth, especially on exposed slopes, even to a depth
of 3 or 4 feet. The wind, prevailingly from some southerly direction
averages about 96 miles a day just above the vegetation on south
slopes. The air temperatures, which may reach 90
0
to 100
0
F., show
a mean daily range varying from 25
0
F. in April and May to 38
0
F.
in July and August. The mean daily soil temperatures at a depth of
3 inches vary from 40
0
to 50
0
F. in May to from 66
0
to 92
0
F. in August
depending upon the slope. The humidity, normally low, frequentl;
falls to 10 or 15 per cent during afternoons. Theaveragedailyevapora-
tion throughout the growing season is about 29 c.c. on southerly slopes
and 20 c.c. on northerly ones. ,
Root penetration and the amount of branching were found in both
the prairies of Washington and those of Nebraska to be profoundly
affected by hard soil. Both the root length and the amount of branch-
ing of a prairie species growing under the two conditions of soil texture
were found to be more pronounced in the less compact soil. Plants
growing in the mellow loess soil of eastern Nebraska are deeper-rooted
than those growing in glacial drift soils. 'Aeration may be an impor-
tant factor in effecting these modifications. Roots show a marked
increase in their output of branches upon leaving compact Boil and
entering earthworm burrows or small crevices. . .:.;:,
The root systems of 28 important plains species were examined near
Colorado Springs, Colorado. They were found to have deeply pene-
trating and widely spreading roots. The lateral spreading of roots in
the surface soil at depths of from 2 to 12 inches is much more prO-
nounced than in either of the preceding prairie communities. Only
four plants, including two cacti, had their root systems confined almost
entirely to the surface 2 feet of soil. One-third of the plains species
penetrated to depths greater than 2 feet but seldom beyond 5 feet.
This group includes such dominant grasses as Bouteloua graCilis and
Aristida purpurea. Nearly 60 per cent of the plains species had roots
extending to depths greater than 5 feet. The roots of several plants
reached depths of 10 to 12 feet.. Although as a community plains
plants are deeper-rooted than those of the prairies, no roots were found
at a depth greater than 13 feet, as was the case with several'prairie
species in Nebraska. It is probable that water never penetrates more
deeply than 13 feet in the plains. . '.
Like the plants of prairies and sandhills, the roots of plains species
show a wide range in type from those with the tap as the principal
feature or with the laterals placed near the surface and especially well
branched, to roots with both tap and laterals well developed. How-
SUMMARY. 125
er the latter or generalized type is the most common. Notwith-
::andmg these individual each community, viewed. as a
whole, has as its own root habIt the one best fitted to the partIcular
environment. . .. .
These plains specIes grow under a mean annual preCIpItatIOn of
about 15 inches, 12 inches of which fall during the growing season.
The soil is a sandy loam so compacted that ordinarily it can be removed
only with a of the of run-off .
after rains is high. MOIsture determInatIOns durmg 1918 show that
the soil to a depth of 7 feet is rather uniformly dry and that at certain
periods during the summer no water is available for plant growth to
a depth of 5 feet. The wind movement over the plants averages about
120 miles per day. The daily fluctuation of temperature among the
plants is usually about 35 to 40 F., the air reaching a maximum of
90 to 95 F. Soil temperatures among the roots at a depth of 4
inches ranged daily from 60 or 70 F. to 90 or 95 F. The average
daily evaporation throughout the growing-season is about 48 c.c.
The well-developed system of shallow, widely spreading laterals is
undoubtedly a response to the moisture in the surface soils resulting
from frequent light summer showers. These surface roots are especially
well developed in cacti. Those of Yucca are remarkable for their great
lateral extent, often reaching a distance of more than 30 feet on every
side of the plant. Bisects show that root competition among plains
plants, especially in the first 2 feet of soil, is very severe.
The root habits of 19 important sandhill species were studied in
eastern Colorado, about 40 miles southeast of Colorado Springs; 8 of
these have roots which are entirely or nearly confined to the surface 2
feet of soil. The deeper-rooted species, none of which were traced
below 11 feet because of the caving sand, practically all show a striking
profusion of long, widely spreading laterals in this surface-soil stratum.
The lower parts of deeply penetrating roots are often much more
poorly branched. Several deep-rooted plains species under sandhill
conditions develop only shallow roots.
General climatic conditions are almost identical with those described
for the plains; hence differences in root habit must be attributed to
edaphic causes. Precipitated moisture is readily absorbed and there
is practically no run-off, even in the heaviest showers. During dry
weather the surface sand forms a very efficient means of retarding
evaporation. Moisture determinations and observations show that
the surface 2 or 3 feet of soil often is moister than the deeper strata,
and thus offers a logical explanation for the typically shallow but
widely spreading sandhill root habit. Several plants have in addition
deeply penetrating roots; hence the deeper soils must receive moisture,
perhaps during wet phases of the climatic cycle. I pomrea leptophylla
has the most extensive root development of any species examined in
the eight communities studied.
- 126 THE ECOLOGICAL RELATIONS OF ROOTS.
In the Rocky Mountains, adjoining the plains at Colorado Springs
the root systems of plants were studied in the gravel-slide, half-gravel:
slide, and forest communities respectively. These communities form.
a developmental series. All plants of the gravel-slide community are
similar in possessing root systems adapted to secure moisture and
nutrients from the surface foot, and few extend beyond a depth of 18
to 24 inches. They have a reIruP'kably wide lateral spread, combined
with a most profuse system of branching.
The surface soil consists of coarse, angular rock fragments,
are from over an inch to only a few millimeters in diameter. Below
this lie a few inches of coarse gravel and sand which gradually give
way, at a depth of 1 to 2.5 feet, to poorly disintegrated granite. The
precipitation falls largely as frequent summer showers which seldom
wet the soil below 18 to 24 inches. The surface efficiently,
both in preventing run-ofI and in retarding evaporation. Moisture
determinations show that the soil moisture, while at no time high, is
rather equally distributed throughout the surface 18 inches of soil.
Plants are so sparsely spaced that relatively little water is removed by
transpiration. Wmd movement averaged 103 miles daily. The aver-
age daily evaporation throughout the summer was 34 c.c. The
Iilunity root habit is clearly a response to the environment. The
development of strong laterals, characteristically up the slope, seems
to be due in part to a response to the soil slipping down the mountains.
Root competition on the gravel-slide is not at all severe. -y
. An examination of roots in the half-gravel-slide revealed quite a dif-
ferent type. In addition to the shallow, widely spreading root habit,
most species had supplementary deep roots which penetrated from
2 to 4 feet. The soil of the former gravel-slide has now become deeper
and richer in humus and much more densely populated. More than
half of the surface is occupied by plants, dominant among which are
certain grasses, while all of the soil to a depth of 3 feet is occupied by
roots. The surface gravel has become more disintegrated and com-
pacted and is less efficient in preventing evaporation. Because of
competition for light, and especially for water, most gravel-slide
plants have disappeared. Although wind movement is somewhat less
and evaporation 'considerably lower than in the former barer habitat,
the water-loss by transpiration from the denser plant population is
higher. During periods of drought plants must rely largely upon
the water in the deeper soil. More favorable deep-soil conditions,
together with intense shallow-root competition, are factors largely
determining root depth._ ,,>
Through a transition chaparral stage, the half-gravel-slide gives
way to forest. Pinus ponderosa, Pseudotsuga mucronata, or Picea engel-
manni is the dominant, depending upon the amount of humus and
moisture. The 19 herbs and shrubs examined on the forest floor were
SUMMARY. 127
relatively shallow-rooted. Almost without exception the major por-
tion of the absorbing system lies within the surface 18 inches. A few
relicts from earlier successional stages had roots which penetrated
somewhat deeper. Many of the roots of tree dominants were found
to be superficial. /
The water-content of the soil offers a logical explanation for this
community root habit. The duff retards run-off as well as evapora-
tion. The trees and shrubs reduce wind movement, lower the tem-
perature and evaporation, and consequently greatly reduce water-
losses by transpiration. Series of moisture determinations show that
the surface 6 inches of soil usually have more available water than the
underlying 6 inches, while the third foot is usually much drier than the
second. This may be accounted for by the frequent summer showers.
Not only are the herbs and shrubs of the forest :Boor shallow-rooted,
but also as a rule the root systems are less extensive in proportion to
the transpiring area than is the case with the half-gravel-slide and the
gravel-slide plants.
The root systems of several species growing in two or more dis-
tinctly different habitats were studied. It was found that in most
cases the root type conformed to the community root habit in which the
species was excavated. Thus profound differences, both in root depth
and position and number of branches, were determined for most
ecads. In general root position conformed strikingly with the distribu-
tion of soil moisture. A few stable species were found which showed
little or no variation of the root system when grown under different
environmental conditions.
The general characters of the root systems of a species are often as
marked and distinctive as are the above-ground vegetative characters.
But the root systems of different species of the same genus, while often
somewhat similar, may be of entirely different types.
A knowledge of the position and competition of roots is indispen-
sable in explaining the phenomena of succession. Since root position
so clearly re:Bects the moisture conditions of the soil, especially when
interpreted in its community relations, a study of the root habits of
plants greatly increases our knowledge of the value of various plants
in indicating lands of agricultural or non-agricultural value. A knowl-
edge of root habits is of further value in solving the problems of the
competition of range species and the improvement of the range.
Finally, root systems indicate the distribution of soil moisture in
various habitats, and thus should aid the forester in selecting sites for
reforestation or afforestation.
BIBLIOGRAPHY.
ALWAY, F. J., ~ al . 1916. The loess soils of the Nebraska portion of the transition region.
Soil SClence 1: Nos.'3, 4, 5. '
BATES, C. G. 1910.' Experiments in sandhill planting. Proc. Soc. Am. For. 5: 59.
CANNON, W. A. 1911. The root systems of desert plants. Carnegie Inst . Wash. Pub. No. 131.
1912. Some features of the root systems of desert plants. Pop. Sci. Mon. 41: 90.
--, -. 1913. Botanical features of the Algerian Sahara. Carnegie Inst. Wash. Pub. No. 178.
---. 1918. The evaluation of the soil-temperature factor in root growth: Plant World
21:64.
---and E. E. FREE. 1917. The ecological significance of soil aeration. Science 45: 178.
CLEMENTS, E. S. 1905. The relation of leaf structure to physical factors. Trans. Am. Mia.
Soc. 26: 19. '
CLEMENTS, F. E. 1904. Formation and succession herbaria. Univ. Nebr. Studies 4: 329.
---. 1905. Research methods in ecology. '
---. 1916. Plant succession. Carnegie Inst. Wash. Pub. No. 242.
KORSTIAN, C. F. 1917. The indicator significance of native vegetation in the determination
of forest sites. Plant World 20: 267.
LoVELAND, G. A. 1912. Summary of climatological data for the United States, Sec. 37, South.
em Nebraska.,
MARKLE, M. S. 1917. Root systems of certain desert plants. Bot. Gu. 64: 177. . ,
POOL, R. J. 1914. A study'of the vtlgetation of the sandhills of Nebraska. Minn. Bot. Studies
4: 189. ' ,
POUND, R., and F. E. CLEMENTS., 1900. The phytogeography of Nebraska.:
PRESTON, C. E. 1900. Observations on the root systems of certain cactacem. Bot: Gu.
30: 348.
ROBBINS, W. W. 1917. The botany of crop plants.
SAMPSON, A. W. 1914 Natural revegetation of range lands based upon growth requirements
and life history of the vegetation. Jour. Agr. Res. 3: 39. ,
---. 1917., Important range plants; their life history and forage value. U. S. Dept.
Agr. Bull. 545. ' ,
SHANTS, H. L. 1911. Natural vegetation as an indicator of the capabilities of land for crop
production in the'Great Plains area. U. S. Dept. Agr., Bur. Pl. Ind. Bull. 201.
TEN EYCK, A. M. 1904. The roots of plants. Kans. Agr. Exp. Sta. Bull. 127. '.
THORNBER, J. J. 1901. The prairie-grass formation in region I. Rep .. Bot. Surv. Nebr. 5:29.
WEAVER, J. E. 1915. A study of the root systems of prairie plants of southeastern Washington.
Plant o ~ l d 14: 227. . "
---. 1917. A study of the vegetation of southeastern Washington and adjacent Idaho.
Univ:Nebr. Studies 17: 1.
---and A. F. THIEL. 1917. Ecological studies in the tension zone between prairie and
wOodland. Rep. Bot.Surv. Nebr., n. s. 1: 1.
WEAVE.R
PLATE. 1
A. One end of a trench used in excavating root systems.
i r ~ l i . ~ tn showing the long rhizomes and shallow roots.
PLATE 2
.\ .. \n eastern Nebraska prairie in June. Erigeron ramosus and Meriolix serrulata in the
foreground and bushy Psoralea tenuifiora in the background.
II . . -\n area dominated by Aurostis hiemalis with Allium mutabile, Achillea mille folium, and
Stipa spartea.
A
B
WEAVER PLATE 3
A
B
c
A. Ancbopogon furcatus.
B . . 4.ndropogon scoparius.
c. Andropogon nutans.
.\
WEAVER
, ' 1/
:
';' ..
I
\
\
I
,
J
I
A. Panicmlt virgaturn, showing rhizomes, course roots, and complete single root.
n. Sl ipa spartea.
PLATE 4
B
PLATE 5
WEAVER
PLATE 6
A. Sporobolus longifolius.
B. Bouteloua gracilis, excavated near the quadrat shown in figure 1.
,'. [ AVER
PLATE 7
.\
A. Silphium laciniatum, roots partially excavated.
B. A1Ilorpha canescens, showing wide lateral spread.
PLATE 7
,\
A. Silphium laciniatum, roots partially excavated.
B. Amorpha canescens, showing wide lateral spread.
A
A. Astragalus crassicarpus, mature root system.
[
'J/ --\j': "
B ~ ... < ...
t '.j.". :" ... ' .rr,' I
I
. .
\
. )
' -""
,.(\ ,
)",
.. ) ..
13. Astragalus crassicarpus,
showing root of young plant.
c
~
Ii!
<
'"
:n
l \ ) \ ~ )
~
( \ t
)
i
~ ,
) i "0
. I ~
( . ~
<\ , IT1
" .,
.. -- , ... _,._-..... .. - ,.,.
C. Baptisia bracteata.
WEAVER
PLATE 10
.' {
A
B
\
)
\
A. Psoralea tenuijlora, the tap-root decayed.
B. Psoralea argophylla, showing entire root in center.
)wEAVER
PLATE 11
.A B c
A. Brauneria pallida. B. Lygodesmia juncea, in two sections. C. Lespedeza capitata.
.\
B
A. Ceanothus ovatus; root of a thirteen-year-old plant; a' is a continuation of a.
B. Amorpha canescens; a' is a continuation of a.
WEAVER
PLATE 13
A
A, Symphoricarpos l'u/garis, showing fine network in surface soil.
B, Rhizomes and runners of Symphoricarpos vulgaris; the horizontal line IS the
ground-line,
WEAVER
PLATE 14
(
A
B
A. Rhus glabra, a portion of the root network with ascending rootlet a.
B. Rh liS glabra, with ascending rootlets.
WEAVER
PLATE 15
.A
B
\
I
\
\;
i
\
! J
}\
,
\
.\
c
A. Corylus americana, the roots shown in two sections.
B. Rosa arkansana, the roots shown in two sections.
C. Corylus americana, rhizomes and roots.
WEAVER PLATE 16
A
B
A. Rosa arkansana, showing method of propagation.
B. Rhus glabra invading subclimax prairie.
WEAVER
'.
I
;;11
. .
A. Yucca glauca, showing the multicipital stems and rhizome habit.
B. Prairie of southeastern Washington.
", I'
PLATE 17
A
C. Meter quadrat in prau'ie, showing Balsamorhiza, Festuca, Lithospermtt1n, and Hieracium.
B
c
A
WEAVER
PLATE 18
A. The plnins association neal' Colorado Springs, showing Arislida purpurea
bunches in Bouleloua gracilis t urf.
B. Psoralea lenuijlora in two sections. C. Yucca glauca.
______ -r ____ ________ r-__ == __ -' ____ __ -. ________ -r ____
r
PLATE 19
---"
o
::-:---..

"'"
]
<:l
<>
' .-
/'

.,
oS

.,

WEAVER
PLATE 20
t
A
B
A. Short-grass plains, showing Boule!ou2 gracilis and Opunli2 polyacanlha.
B. General view of the sandhill community.
WEAVER PLATE 21
B
A. A sandhill community, showing Redfieldia, Petalostemon villosus, Psoralea lanceolata
and Chrysospis villosa, with a socies of Eriogonum microthecum at the left.
B. Roots and rhizomes of Calamovilfa longifolia.
A
PLATE 22
\
,)
WEAVER
.' LATE 23 .
': ..
A
B
A. Consocies of Aletes acaulis on the gravel-slide, with Krynitzkia virgata in the foreground.
B. Quadrat on the gravel-slide, showing detail of surface.
W"AIiR
\
c
A. Aletes acaulis.
B. of fine rootlets of Aleles.
D
c. Smilacina stellata.
D. Thlaspi alpestre.
E. 24
B
WEAVER
A
)
t
\
\
1\ '
" )
','
'l
i ,
i
":, ..
A. Half-gravel-slide, showing EIUIIl1ls Iriticoides and the large bare intervals.
B. Elymus triticoides. C. Rubus deliciosus.
Pl.ATE 26
c
WEAVER PLATE 26
A
c
B
A. Kreleria cristata. B. Calamagrostis purpurascens. c. Polentilla argula glandulosa.
WEAVER PLATE 27
A. Picea engelmanni consociation, showi ng the forest floor.
B. Quadrat in the same spruce forest, showing Haplopappus parryi, Fragaria virginiana,
Thaliclrum fendleri, etc.
WEAVER
PLATE 28
...
A
(",
A. Arctostaphylos lwa-llTsi, showing a portion of the root system.
B. Ribes lacustre, seven years old.
A. Root systems of ecads of Smilacina stellata: a, gravel-slide;
b, spruce forest.
B
B. Bouteloua gracilis from the sandhills.
c
..
(
(
C. Abronia jragrans
from the plains.
~
~
<
'" :D
..,
,...
~
' -i
'"
! ~ : -
WEAVER
. ..-.
f ~
i
.r'.-'
, - .
A. 0 puluster opulijolillS, tIYeh-e years old.
B. CI!1'ysopsis villosa from the sandhills.
PLATt. o.J.
I
-L
A
B

Ecological Relations of Roots

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seoparius.-This grass is figured by Shantz (1911: 56) as

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